Sex on Six Legs: Lessons on Life, Love, and Language from the Insect World - Marlene Zuk (2011)


Introduction: Life on Six Legs

Darwin, C. 2001. The Voyage of the Beagle. Reprint. New York: Modem Library.

Dawkins, R. 2005. Introduction: The illusion of design. Natural History, November.

Dethier, V G. 1964. Microscopic brains. Science 143: 1138–1145.

———. 1981. Fly, rat, and man: The continuing quest for an understanding of behavior. Proceedings of the American Philosophical Society 125: 460–466.

Gal, R., and F. Libersat. 2010. A wasp manipulates neuronal activity in the sub-esophageal ganglion to decrease the drive for walking in its cockroach prey. PLoS ONE 5: E10019.

Gallai, N., J. M. Salles, J. Settele, and B. E. Vaissiere. Economic valuation of the vulnerability of world agriculture confronted with pollinator decline. Ecological Economics 68: 810–821.

Hoyt, E., and T. Schultz, eds. 1999. Insect Lives. New York: John Wiley and Sons.

Losey, J. E., and M. Vaughan. 2006. The economic value of ecological services provided by insects. BioScience 56: 311–323.

Ratnieks, F. L. W. 2006. Can humans learn from insect societies? Nova Acta Leopoldina NF 93: 97–116.

Siebert, C. 2009. Something wild. New York Times, March 5.

Vosshall, L. B. 2007. Into the mind of a fly. Nature 450: 193–197.

Wallechinsky, D., I. Wallace, and A. Wallace. 1977. The Book of Lists. New York: William Morrow and Co.

Zimmer, C. 2010. A wasp finds the seat of the cockroach soul. Discover Blogs, The Loom. April 20. Available at

1. If You're So Smart, Why Aren't You Rich?

Burger, J. M. S., M. Kolss, J. Pont, and T. J. Kawecki. 2008. Learning ability and longevity: A symmetrical evolutionary trade-off in Drosophila. Evolution 62: 1294–1304.

Chanda, S., and E. Caulton. 1999. David Douglas Cunninghan (1843–1914): A biographical profile. Aerobiologia 15: 255–258.

Chittka, L., and E. Leadbeater. 2005. Social learning: Public information in insects. Current Biology 15: R869-R871.

Clare, S. 2006. Honeybees make plans. Journal of Experimental Biology 209: ii.

Collett, T. S. 2008. Insect behaviour: Learning for the future. Current Biology 18: R131-R134.

Csibra, G. 2007. Teachers in the wild. Trends in Cognitive Sciences 11: 95–96.

Cunningham, D. D. 1907. The Plagues and Pleasures of Life in Bengal. London: John Murray.

Dacke, M., and M. V. Srinivasan. 2008. Evidence for counting in insects. Animal Cognition 11: 683–689.

D'Ettorre, P. 2007. Evolution of sociality: You are what you learn. Current Biology 17: R766-R768.

Dukas, R. 2008. Evolutionary biology of insect learning. Annual Review of Entomology 53: 145–160.

Dukas, R., C. Clark, and K. Abbott. 2006. Courtship strategies of male insects: When is learning advantageous? Animal Behaviour 72: 1395–1404.

Dyer, A. G., C. Neumeyer, and L. Chittka. 2005. Honeybee (Apis mellifera) vision can discriminate between and recognise images of human faces. Journal of Experimental Biology 208: 4709–4714.

Dyer, A. G., M. G. P. Rosa, and D. H. Reser. 2008. Honeybees can recognise images of complex natural scenes for use as potential landmarks. Journal of Experimental Biology 211: 1180–1186.

Fabre, J. H. 1981. The Insect World of J. Henri Fabre. Reprint. New York: Harper Colophon.

Fisher, O. 1911. Insect intelligence. Nature 86: 144.

Kennerknecht, I., N. Pluempe, and B. Welling. 2008. Congenital prosopagnosia—A common hereditary cognitive dysfunction in humans. Frontiers in Bioscience 13: 3150–3158.

Kosmos, H. 2008. Through the eyes of a bee. Interview with Adrian Dyer. Humboldt Kosmos. Available at

Leadbeater, E., and L. Chittka. 2007. Social learning in insects—From miniature brains to consensus building. Current Biology 17: R703-R713.

Leadbeater, E., N. E. Raine, and L. Chittka. Social learning: Ants and the meaning of teaching. Current Biology 16: R323-R325.

Mery, F., A. T. Belay, A. K.-C. So, M. B. Sokolowski, and T. J. Kawecki. 2007. Natural polymorphism affecting learning and memory in Drosophila. Proceedings of the National Academy of Sciences USA 104: 13051–13055.

Mery, F., and T.J.Kawecki. 2004. The effect of learning on experimental evolution of resource preference in Drosophila melanogaster. Evolution 58: 757–767.

———. 2005. A cost of long-term memory in Drosophila. Science 308: 1148.

Nowbahari, E., A. Scohier, J.-L. Durand, and K. L. Hollis. 2009. Ants, Catagly-phis cursor, use precisely directed rescue behavior to free entrapped relatives. PLoS ONE 4: E6573.

Paenke, I., B. Sendhoff, and T.J.Kawecki. 2007. Influence of plasticity and learning on evolution under directional selection. American Naturalist 170: E47-E58.

Richardson, T.O., P. A. Sieeman, J.M.McNamara, A. I. Houston, and N. R. Franks. 2007. Teaching with evaluation in ants. Current Biology 17: 1520–1526.

Sitaraman, D., M. Zars, H. LaFerriere, Y.-C. Chen, A. Sable-Smith, T. Kita-moto, G. E. Rottinghaus, and T. Zars. 2008. Serotonin is necessary for place memory in Drosophila. Proceedings of the National Academy of Sciences USA105: 5579–5584.

Thornton, A. 2008. Variation in contributions to teaching by meerkats. Proceedings of the Royal Society of London B 275: 1745–1751.

Thornton, A.,N. J.Raihani, and A. N.Radford. 2007. Teachers in the wild: Some clarification. Trends in Cognitive Sciences 11: 272–273.

Tomchik, S. M., and R. Davis. 2008. Out of sight, but not out of mind. Nature 453: 1192–1194.

Wessnitzer, J., M. Mangan, and B. Webb. 2008. Place memory in crickets. Proceedings of the Royal Society of London B 275: 915–921.

Zhang, S., S. Schwarz, M. Pahl, H. Zhu, and J. Tautz. 2006. Honeybee memory: A honeybee knows what to do and when. Journal of Experimental Biology 209: 4420–4428.

Zimmer, C. 2008. Lots of animals learn, but smarter isn't better. New York Times, May 6.

2. Six Legs and a Genome

Birney, E. 2007. Come fly with us. Nature 450: 184–185.

Brenner, S. 1996. Interview: The world of genome projects. BioEssays 12: 1039–1042.

Chadee, D. D., P. Kittayapong, A. C. Morrison, and W. J. Tabachnick. 2007. A breakthrough for global public health. Science 316: 1703–1704.

Check, E. 2006. From hive minds to humans. Nature 443: 893.

Cusson, M. 2008. The molecular biology toolbox and its use in basic and applied insect science. BioScience 58: 691–700.

Evans, J. D., and D. Gundersen-Rindal. 2003. Beenomes to Bombyx: Future directions in applied insect genomics. Genome Biology 4: 107.

Flannery, M. C. 2007. One genome, one piece of the puzzle. American Biology Teacher 69: 109–112.

———. 2008. Insects by the numbers. American Biology Teacher 70: 426–429.

Gregory, T. R. 2005. Synergy between sequence and size in large-scale genomics. Nature Reviews Genetics 6: 699–708.

———. 2005. Genome size evolution in animals. In The Evolution of the Genome, ed. T. R. Gregory. New York: Elsevier.

Gregory, T. R., and J. S. Johnston. 2008. Genome size diversity in the family Drosophilidae. Heredity 101: 228–238.

Gunter, C. 2007. Genomics on the fly. Nature Reviews Genetics 8: 904.

Jenner, R. A., and M. A. Wills. 2007. The choice of model organisms in evo-devo. Nature Reviews Genetics 8: 311–319.

Koshikawa, S., S. Miyazaki, R. Cornette, T. Matsumoto, and T. Miura. 2008.
Genome size of termites (Insecta, Dictyoptera, Isoptera) and wood roaches (Insecta, Dictyoptera, Cryptocercidae). Naturwissenschaften 95: 859–867.

Ledford, H. 2007. Attack of the genomes. Nature 450: 142–143.

Maderspacher, F. 2008. Genomics: An inordinate fondness for beetles. Current Biology 18: R466.

Myrmecos blog. 2009. Which ants should we target for genome sequencing? January 15. Available at

National Human Genome Research Institute. 2010. NHGRI website. Available at

Pennisi, E. 2007. Fruit fly blitz shows the power of comparative genomics. Science 318: 903.

Ponting, C. P. 2008. The functional repertoires of metazoan genomes. Nature Reviews Genetics 9: 689–698.

Robinson, G. E., and Y. Ben-Shahar. 2002. Social behavior and comparative genomics: New genes or new gene regulation? Genes, Brain, and Behavior 1: 197–203.

Smith, C. R., A. L. Toth, A. V. Suarez, and G. E. Robinson. 2008. Genetic and genomic analyses of the division of labour in insect societies. Nature Reviews Genetics 9: 735–748.

Thompson, G. J., H. Yockey, J. Lim, and B. P. Oldroyd. 2007. Experimental manipulation of ovary activation and gene expression in honey bee (Apis mellifera) queens and workers: Testing hypotheses of reproductive regulation. Journal of Experimental Zoology 307A: 600–610.

Toth, A. L., and G. E. Robinson. 2007. Evo-devo and the evolution of social behavior. Trends in Genetics 23: 334–341.

Toth, A. L., K. Varala, T. C. Newman, F. E. Miguez, S. K. Hutchison, D. A. Willoughby, J. F. Simons, M. Egholm, J. H. Hunt, M. E. Hudson, and G. E. Robinson. 2007. Wasp gene expression supports an evolutionary link between maternal behavior and eusociality. Science 318: 441–444.

Tribolium Genome Sequencing Consortium. 2008. The genome of the model beetle and pest Tribolium castaneum. Nature 452: 949–955.

Tsutsui, N. D., A. V Suarez, J. C. Spagna, and J. S. Johnston. 2008. The evolution of genome size in ants. BMC Evolutionary Biology 8: 64.

Wade, N. 2000. Scientist at work: Sydney Brenner. New York Times, March 7.

Waterhouse, R. M., S. Wyder, and E. M. Zdobnov. 2008. The Aedes aegypti genome: A comparative perspective. Insect Molecular Biology 17: 1–8.

Whitfield, J. 2007. Who's the queen? Ask the genes. Science 318: 910–911.

Wilson, E. O. 2006. How to make a social insect. Nature 443: 919–920.

Zagorski, N. 2006. Profile of Gene E. Robinson. Proceedings of the National Academy of Sciences USA 103: 16065–16067.

Zdobnov, E. M., and P. Bork. 2006. Quantification of insect genome divergence. Trends in Genetics 23: 16–20.

3. The Inner Lives of Wasps

Bell, A. M. 2007. Animal personalities. Nature 447: 539–540.

Biro, P. A., and J. A. Stamps. 2008. Are animal personality traits linked to life-history productivity? Trends in Ecology and Evolution 23: 361–368.

Cervo, R., L. Dapporto, L. Beani, J. E. Strassmann, and'S. Turillazzi. 2008. On status badges and quality signals in the paper wasp Polistes dominulus: Body size, facial colour patterns and hierarchical rank. Proceedings of the Royal Society of London B 275: 1189–1196.

Darwin, C. 2009. The Expression of the Emotions in Man and Animals. Reprint of 1872 edition. New York: Penguin Classics.

Dethier, V. G. 1964. Microscopic brains. Science 143: 1138–1145.

D'Ettorre, P., and J. Heinze. 2005. Individual recognition in ant queens. Current Biology 15: 2170–2174.

Dreier, S., J. S. van Zweden, and P. D'Ettorre. 2007. Long-term memory of individual identity in ant queens. Biology Letters 3: 459–462.

Gosling, S. D. 2001. From mice to men: What can we learn about personality from animal research? Psychological Bulletin 127: 45–86.

Gosling, S. D., and S. Vazire. 2002. Are we barking up the right tree? Evaluating a comparative approach to personality. Journal of Research in Personality 36: 607–614.

Griffin, D. R. 1984. Animal Thinking. Cambridge, MA: Harvard University Press.

———. 2001. Animal Minds. Chicago: University of Chicago Press.

Gronenberg, W., L. E. Ash, and E. A. Tibbetts. 2008. Correlation between facial pattern recognition and brain composition in paper wasps. Brain, Behavior, and Evolution 71: 1–14.

Higgins, L. A., K. M. Jones, and M. L. Wayne. 2005. Quantitative genetics of natural variation of behavior in Drosophila melanogaster: The possible role of the social environment on creating persistent patterns of group activity. Evolution 59: 1529–1539.

Keller, E. F. 1983. A Feeling for the Organism. New York: W H. Freeman and Co.

Koolhaas, J. M. 2008. Coping style and immunity in animals: Making sense of individual variation. Brain, Behavior, and Immunity 22: 662–667.

Kortet, R., and A. Hedrick. 2007. A behavioural syndrome in the field cricket Gryllus integer: Intrasexual aggression is correlated with activity in a novel environment. Biological Journal of the Linnean Society 91: 475–482.

Mehta, P. H., and S. D. Gosling. 2008. Bridging human and animal research: A comparative approach to studies of personality and health. Brain, Behavior, and Immunity 22: 651–661.

Nemiroff, L., and E. Despland. 2007. Consistent individual differences in the foraging behaviour of forest tent caterpillars (Malacosoma disstria). Canadian Journal of Zoology 85: 1117–1124.

Nettle, D. 2006. The evolution of personality variation in humans and other animals. American Psychologist 61: 622–631.

Øyvind, Ø. 2007. Preface: Plasticity and diversity in behavior and brain function—Important raw material for natural selection? Brain, Behavior, and Evolution 70: 215–217.

Réale, D., S. M. Reader, D. Sol, P. T. McDougall, and N. J. Dingemanse. 2007. Integrating animal temperament within ecology and evolution. Biological Reviews 82: 291–318.

Robins, R. W. 2005. The nature of personality: Genes, culture, and national character. Science 310: 62–63.

Sih, A., A. M. Bell, and J. C. Johnson. 2004. Behavioral syndromes: An integrative overview. Quarterly Review of Biology 79: 241–277.

———. 2004. Behavioral syndromes: An ecological and evolutionary overview. Trends in Ecology and Evolution 19: 372–378.

Sih, A., and J. V. Watters. 2005. The mix matters: Behavioural types and group dynamics in water striders. Behaviour 142: 1417–1431.

Stamps, J. A. 2007. Growth-mortality tradeoffs and "personality traits" in animals. Ecology Letters 10: 355–363.

Tibbetts, E. A. 2002. Visual signals of individual identity in the wasp Polistes fuscatus. Proceedings of the Royal Society of London B 269: 1423–1428.

———. 2004. Complex social behaviour can select for variability in visual features: A case study in Polistes wasps. Proceedings of the Royal Society of London B 271: 1955–1960.

Tibbetts, E. A., and J. Dale. 2004. A socially enforced signal of quality in a paper wasp. Nature 432: 218–222.

———. 2007. Individual recognition: It is good to be different. Trends in Ecology and Evolution 22: 529–537.

Tibbetts, E. A., and R. Lindsay. 2008. Visual signals of status and rival assessment in Polistes dominulus paper wasps. Biology Letters 4: 237–239.

Wilson, D. S., A. B. Clark, K. Coleman, and T. Dearstyne. 1994. Shyness and boldness in humans and other animals. Trends in Ecology and Evolution 9: 442–446.

Wolf, M., G. S. van Doorn, O. Leimar, and F. J. Weissing. 2007. Life-history trade-offs favour the evolution of animal personalities. Nature 447: 581–584.

4. Seinfeld and the Queen

Angier, N. 2007. In Hollywood hives, the males rule. New York Times, November 13.

Brackney, S. 2007. The real life of bees. New York Times, November 9.

Charlat, S., E. A. Hornett, J. H. Fullard, N. Davies, G. K. Roderick, N. Wedell, and G. D. D. Hurst. Extraordinary flux in sex ratio. Science 317: 214.

Cobb, M. 2002. Jan Swammerdam on social insects: A view from the seventeenth century. Insectes Sociaux 49: 92–97.

Crane, E. 1999. The World History of Beekeeping and Honey Hunting. New York: Routledge.

Godfray, H. C. J., and J. H. Werren. 1996. Recent developments in sex ratio studies. Trends in Ecology and Evolution 11: 59–63.

Hamilton, W. D. 1967. Extraordinary sex ratios. Science 156: 477–488.

Swammerdam, J. 2004. Information available at

Trivers, R. L., and H. Hare. 1976. Haplodiploidy and the evolution of the social insect. Science 191: 249–263.

Trivers, R. L., and D. E. Willard. 1973. Natural selection of parental ability to vary the sex ratio of offspring. Science 179: 90–92.

Wilson, B. 2004. The Hive: The Story of the Honeybee and Us. London: John Murray.

Zuk, M. 2002. Sexual Selections: What We Can and Can't Learn about Sex from Animals. Berkeley: University of California Press.

5. Sperm and Eggs on Six Legs

Ben-Ari, E. T. 2000. Choosy females. BioScience 50: 7–12.

Birkhead, T. R. 2000. Defining and demonstrating postcopulatory female choice—Again. Evolution 54: 1057–1060.

Birkhead, T. R., and T. Pizzari. 2002. Postcopulatory sexual selection. Nature Reviews Genetics 3: 262–273.

Bjork, A., Dallai, R., and S. Pitnick. 2007. Adaptive modulation of sperm production rate in Drosophila bifurca, a species with giant sperm. Biology Letters 3: 517–519.

Briceño, R. D., W. G. Eberhard, and A. S. Robinson. 2007. Copulation behaviour of Glossina pallidipes (Diptera: Muscidae) outside and inside the female, with a discussion of genitalic evolution. Bulletin of Entomological Research97: 471–488.

Chapman, T. 2008. The soup in my fly: Evolution, form and function of seminal fluid proteins. PLoS Biology 6: 1379–1382.

Córdoba-Aguilar, A. 2006. Sperm ejection as a possible cryptic female choice mechanism in Odonata (Insecta). Physiological Entomology 31: 146–153.

Eberhard, W. G. 1991. Copulatory courtship and cryptic female choice in insects. Biological Reviews 66: 1–31.

Eberhard, W. G., and C. Cordero. 1995. Sexual selection by cryptic female choice on male seminal products—A new bridge between sexual selection and reproductive physiology. Trends in Ecology and Evolution 10: 493–496.

Engqvist, L. 2007. Nuptial gift consumption influences female remating in a scorpionfly: Male or female control of mating rate? Evolutionary Ecology 21: 49–61.

Fedina, T. Y. 2006. Cryptic female choice during spermatophore transfer in Tribolium castaneum (Coleoptera: Tenebrionidae). Journal of Insect Physiology 53: 93–98.

Holland, B., and W. R. Rice. 1997. Cryptic sexual selection—More control issues. Evolution 51: 321–324.

Holman, L., and R. R. Snook. 2008. A sterile sperm caste protects brother fertile sperm from female-mediated death in Drosophila pseudoobscura. Current Biology 18: 292–296.

Jagadeeshan, S., and R. S. Singh. 2006. A time-sequence functional analysis of mating behaviour and genital coupling in Drosophila: Role of cryptic female choice and male sex-drive in the evolution of male genitalia. Journal of Evolutionary Biology 19: 1058–1070.

Joly, D., C. Bressac, and D. Lachaise. 1995. Disentangling giant sperm. Nature 377: 202.

Kullmann, H., and K. P. Sauer. 2008. Mating tactic dependent sperm transfer rates in Panorpa similis (Mecoptera; Panorpidae): A case of female control? Ecological Entomology 34: 153–157.

LaMunyon, C. W., and T. Eisner. 1993. Postcopulatory sexual selection in an arctiid moth (Utetheisa ornatrix). Proceedings of the National Academy of Sciences USA 90: 4689–4692.

Martin, O., and M. Demont. 2008. Reproductive traits: Evidence for sexually selected sperm. Current Biology 18: R79-R81.

Miller, G. T., and S. Pitnick. 2002. Sperm-female coevolution in Drosophila. Science 298: 1230–1233.

Parker, G. 1970. Sperm competition and its evolutionary consequences in the insects. Biological Reviews 45: 525–567.

Pattarini, J. M., W. T. Starmer, A. Bjork, and S. Pitnick. 2006. Mechanisms underlying the sperm quality advantage in Drosophila melanogaster. Evolution 60: 2064–2080.

Peretti, A., W. G. Eberhard, and R. D. Briceno. 2006. Copulatory dialogue: Female spiders sing during copulation to influence male genitalic movements. Animal Behavior 72: 413–421.

Pitnick, S., G. S. Spicer, and T. A. Markow. 1995. How long is a giant sperm? Nature 375: 109.

Pizzari, T. 2006. Evolution: The paradox of sperm leviathans. Current Biology 16: R462-R464.

Simmons, L. W. 2001. Sperm Competition and Its Evolutionary Consequences in the Insects. Princeton, NJ: Princeton University Press.

———. 2005. The evolution of polyandry: Sperm competition, sperm selection, and offspring viability. Annual Review of Ecology, Evolution, and Systematics 36: 125–146.

Simmons, L. W., and F. Garcia-González. 2008. Evolutionary reduction in testes size and competitive fertilization success in response to the experimental removal of sexual selection in dung beetles. Evolution 62: 2580–2591.

Ward, P. I. 2000. Cryptic female choice in the yellow dung fly Scathophaga stercoraria (L.). Evolution 54: 1680–1686.

Wilson, N., S. C. Tubman, P. E. Eady, and G. W Robertson. 1997. Female genotype affects male success in sperm competition. Proceedings of the Royal Society of London B 264: 1491–1495.

6. So Two Fruit Flies Go into a Bar...

Aldous, P. 2008. Randy flies reveal how booze affects inhibitions. New Scientist, January 3.

Bagemihl, B. 1999. Biological Exuberance: Animal Homosexuality and Natural Diversity. New York: St. Martin's Press.

Baram, M. 2007. If there was a gay-straight switch, would you switch? ABC News, December 14. Available at

Featherstone, D. E. 2010. Laboratory website. Available at

Gillespie, R. G. 1991. Homosexual mating behavior in male Doryonychus raptor (Araneae, Tetragnathidae). Journal of Arachnology 19: 229–230.

Grosjean, Y., M. Grillet, H. Augustin, J. F. Ferveur, and D. E. Featherstone. 2008. A glial amino-acid transporter controls synapse strength and courtship in Drosophila. Nature Neuroscience 11: 54–61.

Harari, A. R., H. J. Brockmann, and P. J. Landolt. 2000. Intrasexual mounting in the beetle Diaprepes abbreviatus (L.). Proceedings of the Royal Society of London B 267: 2071–2079.

Khamsi, R. 2005. Fruitflies tap in to their gay side. Nature News, June 2. Available at

———. 2005. Gay flies lose their nerve. BioEd Online, November 9. Available at

Kim, Y.-K., and L. Ehrman. 1998. Developmental isolation and subsequent
adult behavior of Drosophila paulistorum. IV Courtship. Behavior Genetics 28: 57–65.

Kimura, K., T. Hachiya, M. Koganezawa, T. Tazawa, and D. Yamamoto. 2008. Fruitless and doublesex coordinate to generate male-specific neurons that can initiate courtship. Neuron 59: 759–769.

Kyriacou, C. P. 2005. Sex in fruitflies is fruitless. Nature 436: 334–335.

Lee, H-G., Y.-C. Kim, J. S. Dunning, and K.-A. Han. 2008. Recurring ethanol exposure induces disinhibited courtship in Drosophila. PLoS ONE 3: E139.

Levan, K. E., T. Y. Fedina, and S. M. Lewis. 2008. Testing multiple hypotheses for the maintenance of male homosexual copulatory behaviour in flour beetles. Journal of Evolutionary Biology 22: 60–70.

LeVay, S. 1996. Queer Science: The Use and Abuse of Research into Homosexuality. Boston: MIT Press.

Liu, T., L. Dartevelle, C. Yuan, H. Wei, Y. Wang, J.-F. Ferveur, and A. Guo. 2008. Increased dopamine level enhances male-male courtship in Drosophila. Journal of Neuroscience 28: 5539–5546.

McRobert, S. P., and L. Tompkins. 1988. Two consequences of homosexual courtship performed by Drosophila melanogaster and Drosophila affinis males. Evolution 42: 1093–1097.

Miyamoto, T., and H. Amrein. 2008. Suppression of male courtship by a Drosophila pheromone inhibitor. Nature Neuroscience 11: 874–876.

Owen, J. 2005. Damselfly mating game turns some males gay. National Geographic News, June 21. Available at

Preston-Mafham, K. 2006. Post-mounting courtship and the neutralizing of male competitors through "homosexual" mountings in the fly Hydromyza livens F. (Diptera: Scatophagidae). Journal of Natural History 40: 101–105.

Reinhardt, K., E. Harney, R. Naylor, S. Gorb, and M. T. Siva-Jothy. 2007. Female-limited polymorphism in the copulatory organ of a traumatically inseminating insect. American Naturalist 170: 931–935.

Rono, E., P. G. N. Njagi, M. O. Bashir, and A. Hassanali. 2007. Concentration-dependent parsiomonious releaser roles of gregarious male pheromone of the desert locust, Schistocerca gregaria. Journal of Insect Physiology 54: 162–168.

Serrano, J. M., L. Castro, M. A. Toro, and C. Lopez-Fanjul. 1991. The genetic properties of homosexual copulation behavior in Tribolium castaneum: Diallel analysis. Behavior Genetics 21: 547–558.

Switzer, P. V., P. S. Forsythe, K. Escajeda, and K. C. Kruse. 2004. Effects of environmental and social conditions on homosexual pairing in the Japanese Beetle (Popillia japonica Newman). Journal of Insect Behavior 17: 1–16.

Tennent, W. J. 1987. A note on the apparent lowering of moral standards in the Lepidoptera. Entomologists Record 99: 81–82.

Van Gossum, H., L. De Bruyn, and R. Stoks. 2005. Reversible switches between male-male and male-female mating behaviour by male damselflies. Biology Letters 1: 268–270.

Vosshall, L. B. 2008. Scent of a fly. Neuron 59: 685–689.

Wang, Q., L. Chen, J. Li, and X. Yin. 1996. Mating behavior of Phytoecia rufiventris Gautier (Coleoptera: Cerambycidae). Journal of Insect Behavior 9: 47–60.

7. Parenting and the Rotten Corpse

Beal, C. A., and D. W. Tallamy. 2006. A new record of amphisexual care in an insect with exclusive paternal care: Rhynocoris tristis (Heteroptera: Reduviidae). Journal of Ethology 24: 305–307.

Cocroft, R. 2002. Antipredator defense as a limited resource: Unequal predation risk in broods of an insect with maternal care. Behavioral Ecology 13: 125–133.

Costa, J. T. 2006. The Other Insect Societies. Cambridge, MA: Belknap Press.

Evans, T. A., E. J. Wallis, and M. A. Elgar. 1995. Making a meal of mother. Nature 376: 299.

Godfray, H.C.J. 1995. Evolutionary theory of parent-offspring conflict. Nature 376: 133–138.

———. 2005. Quick guide: Parent-offspring conflict. Current Biology 15: R191.

Goubault, M., D. Scott, and I. C. W. Hardy. 2007. The importance of offspring value: Maternal defence in parasitoid contests. Animal Behaviour 74: 437–446.

Klug, H., and M. B. Bonsall. 2007. When to care for, abandon, or eat your offspring: The evolution of parental care and filial cannibalism. American Naturalist 170: 886–901.

Kölliker, M. 2007. Benefits and costs of earwig (Forficula auricularia) family life. Behavioral Ecology and Sociobiology 61: 1489–1497.

Mas, F., and M. Kolliker. 2008. Maternal care and offspring begging in social insects: Chemical signalling, hormonal regulation and evolution. Animal Behaviour 76: 1121–1131.

Nakahira, T., and S. Kudo. 2008. Maternal care in the burrower bug Adomerus triguttulus: Defensive behavior. Journal of Insect Behavior 21: 306–316.

Ohba, S., K. Hidaka, and M. Sasaki. 2006. Notes on paternal care and sibling cannibalism in the giant water bug, Lethocerus deyrolli (Heteroptera: Belostomatidae). Entomological Science 9: 1–5.

Perry, J. C., and B. D. Roitberg. 2005. Ladybird mothers mitigate offspring starvation risk by laying trophic eggs. Behavioral Ecology and Sociobiology 58: 578–586.

———. 2006. Trophic egg laying: Hypotheses and tests. Oikos 112: 706–714.

Roy, H. E., H. Rudge, L. Goldrick, and D. Hawkins. 2007. Eat or be eaten: Prevalence and impact of egg cannibalism on two-spot ladybirds, Adalia bipunctata. Entomologia Experimentalis et Applicata 125: 33–38.

Santi, F., and S. Maini. 2007. Ladybirds mothers eating their eggs: Is it cannibalism? Bulletin of Insectology 60: 89–91.

Saul-Gershenz, L. S., and J. G. Millar. 2006. Phoretic nest parasites use sexual deception to obtain transport to their host's nest. Proceedings of the National Academy of Sciences USA 103: 14039–14044.

Smiseth, P. T., and H. J. Parker. 2008. Is there a cost to larval begging in the burying beetle Nicrophorus vespilloides? Behavioral Ecology 19: 1111–1115.

Smiseth, P. T., R.J.S. Ward, and A. J. Moore. 2006. Asynchronous hatching in Nicrophorus vespilloides, an insect in which parents provide food for their offspring. Functional Ecology 20: 151–156.

Smith, G., S. T. Trumbo, D. S. Sikes, M. P. Scott, and R. L. Smith. 2007. Host shift by the burying beetle, Nicrophorus pustulatus, a parasitoid of snake eggs. Journal of Evolutionary Biology 20: 2389–2399.

Smith, R. L. 1979. Paternity assurance and altered roles in the mating behaviour of a giant water bug, Abedus herberti (Heteroptera, Belostomatidae). Animal Behaviour 27: 716–725.

Staerkle, M., and M. Kolliker. 2008. Maternal food regurgitation to nymphs in earwigs (Forficula auricularia). Ethology 114: 844–850.

Steiger, S., K. Peschke, W. Francke, and J. K. Muller. 2007. The smell of parents: Breeding status influences cuticular hydrocarbon pattern in the burying beetle Nicrophorus vespilloides. Proceedings of the Royal Society of London B 274: 2211–2220.

Tallamy, D. W. 2005. Egg dumping in insects. Annual Review of Entomology 50: 347–370.

Tallamy, D. W., E. Walsh, and D. C. Peck. 2004. Revisiting parental care in the assassin bug, Atopozeluspallens (Heteroptera, Reduviidae). Journal of Insect Behavior 17: 431–436.

Tallamy, D. W., and T. K. Wood. 1986. Convergence patterns in subsocial insects. Annual Review of Entomology 31: 369–390.

Thomas, L. K., and A. Manica. 2003. Filial cannibalism in an assassin bug. Animal Behaviour 66: 205–210.

Trivers, R. L. 1974. Parent-offspring conflict. American Zoologist 14: 249–264.

Trumbo, S. T. 2006. Infanticide, sexual selection and task specialization in a biparental burying beetle. Animal Behaviour 72: 1159–1167.

———. 2007. Defending young biparentally: Female risk-taking with and without a male in the burying beetle, Nicrophorus pustulatus. Behavioral Ecology and Sociobiology 61: 1717–1723.

Williams, L., III, M. C. Coscaron, P. M. Dellape, and T. M. Roane. 2005. The shield-backed bug, Pachycoris stallii: Description of immature stages, effect of maternal care on nymphs, and notes on life history. Journal of Insect Science 5: 1–13.

Zink, A. G. 2003. Quantifying the costs and benefits of parental care in female treehoppers. Behavioral Ecology 14: 687–693.

8. Pirates at the Picnic

Beebe, W. 1999. The hometown of the army ants. In Insect Lives, ed. E. Hoyt and T. Schultz. Reprint of 1921 edition. New York: John Wiley and Sons.

Beibl, J., R. J. Stuart, J. Heinze, and S. Foitzik. 2005. Six origins of slavery in formicoxenine ants. Insectes Sociaux 52: 291–297.

Bonckaert, W., K. Vuerinckx, J. Billen, R. L. Hammond, L. Keller, and T. Wenseleers. 2008. Worker policing in the German wasp Vespula germanica. Behavioral Ecology 19: 272–278.

Bono, J. M., M. F. Antolin, and J. M. Herbers. 2006. Parasite virulence and host resistance in a slave-making ant community. Evolutionary Ecology Research 8: 1117–1128.

Bono, J. M., E. R. Gordon, M. F. Antolin, and J. M. Herbers. 2006. Raiding activity of an obligate (Polyergus breviceps) and two facultative (Formica puberula and F. gynocrates) slave-making ants. Journal of Insect Behavior 19: 429–446.

Crompton, J. 1954. Ways of the Ant. Boston: Houghton Mifflin Co.

Foitzik, S., C. J. DeHeer, D. N. Hunjan, and J. M. Herbers. 2001. Coevolution in host-parasite systems: Behavioural strategies of slave-making ants and their hosts. Proceedings of the Royal Society of London B 268: 1139–1146.

Gadagkar, R. 2004. Why do honey bee workers destroy each other's eggs? Journal of Bioscience 29: 213–217.

Gloag, R., T. A. Heard, M. Beekman, and B. P. Oldroyd. 2008. Nest defence in a stingless bee: What causes fighting swarms in Trígona carbonaria (Hymenoptera, Meliponini)? Insectes Sociaux 55: 387–391.

Helanterä, H. 2007. How to test an inclusive fitness hypothesis—Worker reproduction and policing as an example. Oikos 116: 1782–1788.

Herbers, J. M. 2006. The loaded language of science. Chronicle of Higher Education 52: B5.

———. 2007. Watch your language! Racially loaded metaphors in scientific research. BioScience 57: 104–105.

Herbers, J. M., and S. Foitzik. 2002. The ecology of slavemaking ants and their hosts in north temperate forests. Ecology 83: 148–163.

Hölldobler, B., and E. O. Wilson. 1990. The Ants. Cambridge, MA: Belknap Press.

———. 1994. Journey to the Ants. Cambridge, MA: Belknap Press.

Hoyt, E., and T. Schultz, eds. 1999. Insect Lives. New York: John Wiley and Sons.

Johnson, C. A., and J. M. Herbers. 2006. Impact of parasite sympatry on the geographic mosaic of coevolution. Ecology 87: 382–394.

Maeterlinck, M. 1930. The Life of the Ant. New York: John Day Co.

Ratnieks, F.L.W., and P. K. Visscher. 1989. Worker policing in the honeybee. Nature 342: 796–797.

Ratnieks, F.L.W, and T. Wenseleers. 2005. Policing insect societies. Science 307: 54–56.

Sleigh, C. 2003. Ant. London: Reaktion Books.

———. 2007. Six Legs Better. Baltimore: Johns Hopkins University Press.

Smith, A. A., and K. L. Haight. 2008. Army ants as research and collection tools. Journal of Insect Science 8: 71–76.

Smith, A. A., B. Hölldober, and J. Liebig. 2009. Cuticular hydrocarbons reliably identify cheaters and allow enforcement of altruism in a social insect. Current Biology 19: 78–81.

Visscher, P. K., and R. Dukas. 1995. Honey bees recognize development of nestmates' ovaries. Animal Behaviour 49: 542–544.

Wenseleers, T., and F. L. W. Ratnieks. 2006. Enforced altruism in insect societies. Nature 444: 50.

Wheeler, W. M., and T. Schneirla. 1934. Raiding and other outstanding phenomena in the behavior of army ants. Proceedings of the National Academy of Sciences USA 20: 316–321.

9. Six-Legged Language

Aleksiev, A. S., B. Longdon, M. J. Christmas, A. B. Sendova-Franks, and N. R. Franks. 2008. Individual and collective choice: Parallel prospecting and mining in ants. Naturwissenschaften 95: 301–305.

Beekman, M., and J. B. Lew. 2008. Foraging in honeybees—When does it pay to dance? Behavioral Ecology 19: 255–262.

Beekman, M., R. S. Gloag, N. Even, W Wattanachaiyingchareon, and B. P. Oldroyd. 2008. Dance precision of Apis florea —Clues to the evolution of the honeybee dance language? Behavioral Ecology and Sociobiology 62: 1259–1265.

Cerdá, X., E. Angulo, and R. Boulay. 2009. Individual and collective foraging decisions: A field study of worker recruitment in the gypsy ant Aphaeno-gaster senilis. Behavioral Ecology and Sociobiology 63: 551–562.

Conradt, L. 2008. Group decisions: How (not) to choose a restaurant with friends. Current Biology 18: R1139–1140.

Conradt, L., and C. List. 2009. Group decisions in humans and animals: A survey. Philosophical Transactions of the Royal Society B 364: 719–742.

Conradt, L., and T. J. Roper. 2005. Consensus decision making in animals. Trends in Ecology and Evolution 20: 449–456.

Couzin, I. D. 2008. Collective cognition in animal groups. Trends in Cognitive Sciences 13: 36–43.

Couzin, I. D., J. Krause, N. R. Franks, and S. A. Levin. 2006. Effective leadership and decision-making in animal groups on the move. Nature 433: 513–516.

Crist, E. 2004. Can an insect speak? The case of the honeybee dance language. Social Studies of Science 34: 7–43.

Detrain, C., and J.-L. Deneubourg. 2008. Collective decision-making and foraging patterns in ants and honeybees. Advances in Insect Physiology 35: 123–173.

Dussutour, A., S. C. Nicolis, E. Despland, and S. J. Simpson. 2008. Individual differences influence collective behaviour in social caterpillars. Animal Behaviour 76: 5–16.

Dussutour, A., S. J. Simpson, E. Despland, and N. Colasurdo. 2007. When the group denies individual nutritional wisdom. Animal Behaviour 74: 931–939.

Dyer, J. R. G., C. C. Ioannou, L. J. Morrell, D. P. Croft, I. D. Couzin, D. A. Waters, and J. Krause. 2008. Consensus decision making in human crowds. Animal Behaviour 75: 461–470.

Franks, N. R., A. Dornhaus, C. S. Best, and E. L. Jones. 2006. Decision making by small and large house-hunting ant colonies: One size fits all. Animal Behaviour 72: 611–616.

Franks, N. R., J. W. Hooper, A. Dornhaus, P. J. Aukett, A. L. Hayward, and S. M. Berghoff. 2007. Reconnaissance and latent learning in ants. Proceedings of the Royal Society B 274: 1505–1509.

Gorman, J. 2006. Mr. Speaker, I'd Like to Do the Waggle. New York Times, May 2.

Hauser, M. D., N. Chomsky, and W. T. Fitch. 2002. The faculty of language: What is it, who has it, and how did it evolve? Science 298: 1569–1579.

Lindauer, M. 1957. Communication in swarm-bees searching for a new home. Nature 179: 63–66.

Maeterlinck, M. 1901. The Life of the Bee. New York: Dodd, Mead and Co.

Marquis, D. 1987. Archy and Mehitabel. Reprint of 1927 edition. New York: Anchor.

Munz, T. 2005. The Bee Battles: Karl von Frisch, Adrian Wenner and the Honey Bee Dance Language Controversy. Journal of the History of Biology 38: 535–570.

Nieh, J. C., L. S. Barreto, F. A. L. Contrera, and V L. Imperatriz-Fonseca. 2004. Olfactory eavesdropping by a competitively foraging stingless bee, Trigona spinipes. Proceedings of the Royal Society of London B 271: 1633–1640.

Passino, K. M., T. D. Seeley, and P. K. Visscher. 2008. Swarm cognition in honey bees. Behavioral Ecology and Sociobiology 62: 401–414.

Pinker, S., and R. Jackendoff. 2005. The faculty of language: What's special about it? Cognition 95: 201–236.

Planqué, R., A. Dornhous, N. R. Franks, T. Kovacs, and J. A. R. Marshall. 2006. Weighing waiting in collective decision-making. Behavioral Ecology and Sociobiology 61: 347–356.

Pollick, A. S., and F. B. M. de Waal. 2007. Ape gestures and language evolution. Proceedings of the National Academy of Sciences USA 104: 8184–8189.

Rittschof, C. C., and T. D. Seeley. 2008. The buzz-run: How honeybees signal "Time to go!" Animal Behaviour 75: 189–197.

Schultz, K. M., K. M. Passino, and T. D. Seeley. 2008. The mechanism of flight guidance in honeybee swarms: Subtle guides or streaker bees? Journal of Experimental Biology 211: 3287–3295.

Seeley, T. D., and P. K. Visscher. 2008. Sensory coding of nest-site value in honeybee swarms. Journal of Experimental Biology 211: 3691–3697.

Seeley, T. D., P. K. Visscher, and K. M. Passino. 2006. Group decision making in honey bee swarms. American Scientist 94: 220–229.

Sherman, G., and P. K. Visscher. 2002. Honeybee colonies achieve fitness through dancing. Nature 419: 920–922.

Skorupski, P., and L. Chittka. 2006. Animal cognition: An insect's sense of time? Current Biology 16: R851-R853.

Smith, E. M., and G. W Otis. 2006. Resolution of a controversy: Functionality of the dance language of the honey bee, Part I. American Bee Journal 3: 242–246.

———. 2006. Resolution of a controversy: Functionality of the dance language of the honey bee, Part II. American Bee Journal 4: 335–340.

Su, S., F. Cai, A. Si, S. Zhang, J. Tautz, and S. Chen. 2008. East learns from West: Asiatic honeybees can understand dance language of European honeybees. PLoS ONE 3: E2365.

Visscher, P. K. 2007. Group decision making in nest-site selection among social insects. Annual Review of Entomology 52: 255–275.

Wenner, A. M. 2002. The elusive honey bee dance "language" hypothesis. Journal of Insect Behavior 15: 859–878.

Wray, A. 2005. The broadening scope of animal communication research. In Language Origins: Perspectives on Evolution, ed. M. Tallerman. Oxford: Oxford University Press.

Wray, M. K., B. A. Klein, H. R. Mattila, and T. D. Seeley. 2008. Honeybees do not reject dances for "implausible" locations: Reconsidering the evidence for cognitive maps in insects. Animal Behaviour 76: 261–269.

Yang, C., P. Belawat, E. Hafen, L. Y. Jan, and Y.-N. Jan. 2008. Drosophila egg-laying site selection as a system to study simple decision-making processes. Science 319: 1679–1683.