Kin Selection and Social Insects Social insects provide the most surprising predictions and satisfying tests of kin selection

Social insects so dominate many terrestrial habitats (Wilson 1990) that they can hardly escape the attention of biologists, but even if they were rare, they would still attract special interest because of the intricate cooperation within their societies. William Morton Wheeler (1911) described the social insect colony as an organism (or as a higher-level organism or superorganism) because of the degree to which individuals appear to operate as a unit that is dedicated to the perpetuation and reproduction of the colony as a whole. The reinvention of the organism at a higher level has occurred at a number of crucial junctures in the history of life (Maynard Smith and Szathmary 1995). For example, the eukaryotic cell arose from several prokaryotic ancestors (Margulis 1970), and multicellular plants, animals, and fungi arose from single-celled ancestors (Buss 1987). Because insect societies are macroscopic, and because they span the entire range from solitary individuals to essentially superorganismal colonies, they offer an accessible model for how such transitions can happen.

[1]  T. Seeley The honey bee colony as a superorganism. , 1989 .

[2]  M. Breed,et al.  Kin discrimination within honey bee (Apis mellifera) colonies: An analysis of the evidence , 1994, Behavioural Processes.

[3]  D. Gordon The organization of work in social insect colonies , 1996, Nature.

[4]  P. Pamilo Evolution of Colony Characteristics in Social Insects. I. Sex Allocation , 1991, The American Naturalist.

[5]  L. Keller,et al.  The role of queen pheromones in social insects: queen control or queen signal? , 1993, Animal Behaviour.

[6]  A. Forsyth Worker Control of Queen Density in Hymenopteran Societies , 1980, The American Naturalist.

[7]  J. Seger,et al.  Kinship and covariance. , 1981, Journal of theoretical biology.

[8]  M. Solignac,et al.  Monoandry and polyandry in bumble bees (Hymenoptera; Bombinae) as evidenced by highly variable microsatellites , 1995, Molecular ecology.

[9]  A. Grafen,et al.  Unmatedness and the Evolution of Eusociality , 1988, The American Naturalist.

[10]  A. Buschinger Regulation of Worker and Queen Formation in Ants with Special Reference to Reproduction and Colony Development , 1990 .

[11]  R. Trivers,et al.  Haplodiploidy and the Evolution of the Social Insects , 2008 .

[12]  L. Packer,et al.  Relatedness and sex ratio in a primitively eusocial halictine bee , 2004, Behavioral Ecology and Sociobiology.

[13]  M. Fondrk,et al.  Paternity analysis of worker honeybees using random amplified polymorphic DNA , 1993, Naturwissenschaften.

[14]  H. Kern Reeve,et al.  Conflict in single-queen hymenopteran societies : the structure of conflict and processes that reduce conflict in advanced eusocial species , 1992 .

[15]  L. Keller,et al.  Indiscriminate altruism: unduly nice parents and siblings. , 1997, Trends in ecology & evolution.

[16]  W. M. Wheeler,et al.  The ant‐colony as an organism , 1911 .

[17]  J. Evans,et al.  Relatedness threshold for the production of female sexuals in colonies of a polygynous ant, Myrmica tahoensis, as revealed by microsatellite DNA analysis. , 1995, Proceedings of the National Academy of Sciences of the United States of America.

[18]  B. Crespi,et al.  Three conditions for the evolution of eusociality: Are they sufficient? , 1994, Insectes Sociaux.

[19]  Richard E. Michod,et al.  The Theory of Kin Selection , 1982 .

[20]  B. Cole Dominance hierarchies in leptothorax ants. , 1981, Science.

[21]  Bernard J. Crespi,et al.  Eusociality in Australian gall thrips , 1992, Nature.

[22]  John Maynard Smith,et al.  The logic of asymmetric contests , 1976, Animal Behaviour.

[23]  J. Boomsma Adaptive colony sex ratios in primitively eusocial bees. , 1991, Trends in ecology & evolution.

[24]  E. Sober,et al.  Reviving the superorganism. , 1989, Journal of theoretical biology.

[25]  A. Grafen,et al.  Colony‐level sex ratio selection in the eusocial Hymenoptera , 1991 .

[26]  L. Hurst,et al.  Genetic Conflicts , 1996, The Quarterly Review of Biology.

[27]  D. Queller Models of kin selection on seed provisioning , 1984, Heredity.

[28]  Stefan Fuchs,et al.  Lower performance in honeybee colonies of uniform paternity , 1994 .

[29]  J. Strassmann,et al.  Maternity assignment and queen replacement in a social wasp , 1995, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[30]  L. Sundström Sex ratio bias, relatedness asymmetry and queen mating frequency in ants , 1994, Nature.

[31]  P. Pamilo Evolution of Colony Characteristics in Social Insects. II. Number of Reproductive Individuals , 1991, The American Naturalist.

[32]  D. Queller,et al.  ESTIMATING RELATEDNESS USING GENETIC MARKERS , 1989, Evolution; international journal of organic evolution.

[33]  Diana E. Wheeler,et al.  Developmental and Physiological Determinants of Caste in Social Hymenoptera: Evolutionary Implications , 1986, The American Naturalist.

[34]  R. Craig Subfertility and the evolution of eusociality by kin selection , 1983 .

[35]  D. Yanega Social plasticity and early-diapausing females in a primitively social bee. , 1988, Proceedings of the National Academy of Sciences of the United States of America.

[36]  R. Jeanne,et al.  The organization of work in Polybia occidentalis: costs and benefits of specialization in a social wasp , 1986, Behavioral Ecology and Sociobiology.

[37]  J. Strassmann,et al.  Age Is More Important Than Size in Determining Dominance Among Workers in the Primitively Eusocial Wasp, Polistes Instabilis , 1988 .

[38]  P. Nonacs Ant Reproductive Strategies and Sex Allocation Theory , 1986, The Quarterly Review of Biology.

[39]  H. Evans Commentary: Extrinsic versus Intrinsic Factors in the Evolution of Insect Sociality , 1977 .

[40]  V. Imperatriz-Fonseca,et al.  Caste Development, Reproductive Strategies, and Control of Fertility in Honey Bees and Stingless Bees , 1990 .

[41]  K. Ross,et al.  Regulation of Reproduction in Eusocial Hymenoptera , 1985 .

[42]  K. Ross,et al.  AN UNUSUAL PATTERN OF GENE FLOW BETWEEN THE TWO SOCIAL FORMS OF THE FIRE ANT SOLENOPSIS INVICTA , 1993, Evolution; international journal of organic evolution.

[43]  L. Keller,et al.  Conditional Manipulation of Sex Ratios by Ant Workers: A Test of Kin Selection Theory , 1996, Science.

[44]  Subfamily recognition and task specialisation in honey bees (Apis mellifera L.) (Hymenoptera : Apidae) , 1994, Behavioral Ecology and Sociobiology.

[45]  W. Hamilton,et al.  Altruism and Related Phenomena, Mainly in Social Insects , 1972 .

[46]  J. Strassmann,et al.  Control of reproduction in social insect colonies: individual and collective relatedness preferences in the paper wasp, Polistes annularis , 1997, Behavioral Ecology and Sociobiology.

[47]  P. Pamilo,et al.  Reproductive competition and conflicts in colonies of the ant Formica sanguinea , 1994, Animal Behaviour.

[48]  J. Evans,et al.  Parentage analyses in ant colonies using simple sequence repeat loci , 1993, Molecular ecology.

[49]  A. Łomnicki,et al.  Multiple mating of queens and the sterility of workers among eusocial hymenoptera , 1987 .

[50]  P. Frumhoff,et al.  Nepotism in the honey bee , 1990, Nature.

[51]  W. Foster,et al.  THE EVOLUTION OF SOLDIERS IN APHIDS , 1996, Biological reviews of the Cambridge Philosophical Society.

[52]  F. Ratnieks Reproductive Harmony via Mutual Policing by Workers in Eusocial Hymenoptera , 1988, The American Naturalist.

[53]  A. Grafen Split sex ratios and the evolutionary origins of eusociality , 1986 .

[54]  Christopher K. Starr,et al.  Sperm competition, kinship, and sociality in the aculeate Hymenoptera , 1984 .

[55]  W. E. Kerr Genetic Determination of Castes in the Genus Melipona. , 1950, Genetics.

[56]  U. Mueller Haplodiploidy and the Evolution of Facultative Sex Ratios in a Primitively Eusocial Bee , 1991, Science.

[57]  A. Bourke,et al.  Split sex ratios in a multiple-queen ant population , 1994, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[58]  J. Simpson,et al.  Eusociality in the beetleAustroplatypus incompertus (Coleoptera: Curculionidae) , 1992, Naturwissenschaften.

[59]  R. Gadagkar Evolution of eusociality: the advantage of assured fitness returns , 1990 .

[60]  J. Strassmann,et al.  A selfish strategy of social insect workers that promotes social cohesion , 1993, Nature.

[61]  Francis L. W. Ratnieks,et al.  Worker policing in the honeybee , 1989, Nature.

[62]  G. B. Pollock Kin selection, kin avoidance and correlated strategies , 2005, Evolutionary Ecology.