J. J. Boomsma, Adaptive colony sex ratios in primitively eusocial bees, Trends Ecol Evol, vol.6, pp.92-95, 1991.

J. J. Boomsma, Sex ratio variation in polygynous ants, Queen number and sociality in insects, pp.86-109, 1993.

J. J. Boomsma and A. Grafen, Intraspecific variation in ant sex ratios and the Trivers-Hare hypothesis, Evolution, vol.44, pp.1026-1034, 1990.

J. J. Boomsma and A. Grafen, Colony-level sex ratio selection in the eusocial Hymenoptera, J Evol Biol, vol.4, pp.383-407, 1991.

J. J. Boomsma, J. Nielsen, L. Sundström, N. J. Oldham, J. Tentschert et al., Informational constraints on optimal sex allocation in ants, Proc Natl Acad Sci, vol.100, pp.8799-8804, 2003.

A. Bourke, Genetics, relatedness and social behaviour in insect societies, Insect evolutionary ecology, pp.1-30, 2005.

A. Bourke and N. R. Franks, Social evolution in ants, 1995.

M. V. Brian, Habitat differences in sexual production by two coexistent ants, J Anim Ecol, vol.48, pp.943-953, 1979.

W. D. Brown and L. Keller, Colony sex ratios vary with queen number but not relatedness asymmetry in the ant Formica exsecta, Proc R Soc B, vol.267, pp.1751-1757, 2000.

, The relatedness asymmetry, measured with 4 microsatellite markers, was 2.66:1 and 3.32:1 for monogyne and polygyne colonies, respectively. These values do not differ significantly, and the magnitude of relatedness asymmetry variation was estimated with the relatedness asymmetry of monogyne colonies in the numerator, Sex investment ratio in monogyne versus polygyne colonies, F 1,22 ¼ 60.13

, Correlation between proportion of females and queen number in polygyne colonies, r s ¼ ÿ0

, Proportion of females produced by monogyne versus polygyne colonies, data from Tables 2-4, excluding queenless colonies, 1-tailed Wilcoxon rank-sum test, vol.369

, Correlation between sex investment ratio and queen number in polygyne colonies, data from Tables 2-4, excluding queenless colonies, Spearman rank correlation test, r s ¼ ÿ0, p.37

, The effect size is conservative because it had to be calculated from the P value threshold. We calculated the relatedness asymmetry as described in Methods, Relatedness among females in female-producing colonies versus relatedness among workers in male-producing colonies, 1-tailed t-test, P , 0.005

, Relatedness among females in female-producing colonies versus relatedness among workers in male-producing colonies, t-test, P , 0.05 for each of the 4 species. Effect sizes are conservative because they had to be calculated from the P value thresholds. We calculated the relatedness asymmetry as described in Methods, using relatedness among queens and relatedness among workers in female-and male-producing nests

, Sex investment ratio in colonies founded by one versus several queens, data from Table 1, Wilcoxon rank-sum test, W ¼ 41.5, P ¼ 0.113. 26 Proportion of haploid eggs in polygyne versus monogyne colonies, v 2 ð4Þ ¼ 2:161. The OH test on these statistics gave P ¼ 0.086. 27 Proportion of ergatoid males in polygyne versus monogyne colonies

, Correlation between proportional investment in females and total sexual productivity, Z ¼ ÿ0, vol.334, p.388

, Behavioral Ecology

A. Buschinger and J. Heinze, Stenamma debile (Hymenoptera, Formicidae): productivity and sex allocation across three years, Insectes Soc, vol.48, pp.110-117, 2001.

G. L. Chan, A. Hingle, and A. Bourke, Sex allocation in a facultatively polygynous ant: between-population and between-colony variation, Behav Ecol, vol.10, pp.409-421, 1999.

M. Chapuisat and L. Keller, Testing kin selection with sex allocation data in eusocial Hymenoptera, Heredity, vol.82, pp.473-478, 1999.

A. B. Clark, Sex ratio and local resource competition in a prosimian primate, Science, vol.201, pp.163-165, 1978.

R. H. Crozier and P. Pamilo, Evolution of social insect colonies: sex allocation and kin selection, 1996.

L. De-menten, S. Cremer, J. Heinze, and A. S. , Primary sex ratio adjustment by ant queens in response to local mate competition, 2005.

, Anim Behav, vol.69, pp.1031-1035

L. De-menten, D. Fournier, C. Brent, L. Passera, E. L. Vargo et al., Dual mechanism of queen influence over sex ratio in the ant Pheidole pallidula, Behav Ecol Sociobiol, vol.58, pp.527-533, 2005.

R. J. Deslippe and R. Savolainen, Sex investment in a social insect: the proximate role of food, Ecology, vol.76, pp.375-382, 1995.

M. J. Duchateau, H. Velthuis, and J. J. Boomsma, Sex ratio variation in the bumblebee Bombus terrestris, Behav Ecol, vol.15, pp.71-82, 2004.

G. W. Elmes, Temporal variation in colony populations of the ant Myrmica sulcinodis. II. Sexual production and sex ratios, J Anim Ecol, vol.56, pp.573-583, 1987.

J. D. Evans, Relatedness threshold for the production of female sexuals in colonies of a polygynous ant, Myrmica tahoensis, as revealed by microsatellite DNA analysis, Proc Natl Acad Sci, vol.92, pp.6514-6517, 1995.

E. J. Fjerdingstad, P. J. Gertsch, and L. Keller, Why do some social insect queens mate with several males? Testing the sex ratio manipulation hypothesis in Lasius niger, Evolution, vol.56, pp.553-562, 2002.

K. R. Foster and F. Ratnieks, The effect of sex allocation biasing on the evolution of worker policing in hymenopteran societies, Am Nat, vol.158, pp.615-623, 2001.

D. Fournier, L. Keller, L. Passera, and A. S. , Colony sex ratios vary with breeding system but not relatedness asymmetry in the facultatively polygynous ant Pheidole pallidula, Evolution, vol.57, pp.1336-1342, 2003.

S. Fuchs and V. Schade, Lower performance in honeybee colonies of uniform paternity, Apidologie, vol.25, pp.155-168, 1994.
URL : https://hal.archives-ouvertes.fr/hal-00891147

A. Gardner and S. A. West, Spite and the scale of competition, J Evol Biol, vol.17, pp.1195-1203, 2004.

A. S. Griffin, B. C. Sheldon, and S. A. West, Cooperative breeders adjust offspring sex ratios to produce helpful helpers, Am Nat, vol.166, pp.628-632, 2005.

W. D. Hamilton, The genetical evolution of social behaviour, J Theor Biol, vol.7, pp.1-52, 1964.

W. D. Hamilton, Extraordinary sex ratios, Science, vol.156, pp.477-488, 1967.
DOI : 10.1126/science.156.3774.477

W. D. Hamilton, Selfish and spitful behaviour in an evolutionary model, Nature, vol.228, pp.1218-1220, 1970.

W. D. Hamilton, Altruism and related phenomena, mainly in social insects, Annu Rev Ecol Syst, vol.3, pp.193-232, 1972.

R. L. Hammond, M. W. Bruford, and A. Bourke, Ant workers selfishly bias sex ratios by manipulating female development, Proc R Soc B, vol.269, pp.173-178, 2002.

M. D. Hastings, D. C. Queller, F. Eischen, and J. E. Strassmann, Kin selection, relatedness, and worker control of reproduction in a large-colony epiponine wasp, Brachygastra mellifica, Behav Ecol, vol.9, pp.573-581, 1998.

J. Heinze, A. Bottcher, and S. Cremer, Production of winged and wingless males in the ant, Cardiocondyla minutior, Insectes Soc, vol.51, pp.275-278, 2004.

J. Heinze, A. Hartmann, and O. Rü, Sex allocation ratios in the facultatively polygynous ant, Leptothorax acervorum, Behav Ecol Sociobiol, vol.50, pp.270-274, 2001.

K. R. Helms, Colony sex ratios, conflict between queens and workers, and apparent queen control in the ant Pheidole desertorum, Evolution, vol.53, pp.1470-1478, 1999.

K. R. Helms, D. Fournier, L. Keller, L. Passera, and A. S. , Colony sex ratios in the facultatively polygynous ant Pheidole pallidula: a reanalysis with new data, Evolution, vol.58, pp.1141-1142, 2004.

J. M. Herbers, Caste-biased polyethism in a mound-building ant species, Am Midl Nat, vol.101, pp.69-75, 1979.

J. M. Herbers, Queen-worker conflict and eusocial evolution in a polygynous ant species, Evolution, vol.38, pp.631-643, 1984.

J. M. Herbers, Reproductive investment and allocation ratios for the ant Leptothorax longispinosus: sorting out the variation, Am Nat, vol.136, pp.178-208, 1990.

E. A. Herre, Optimality, plasticity and selective regime in fig wasp sex ratio, Nature, vol.329, pp.627-629, 1987.

C. R. Hughes, D. C. Queller, J. E. Strassmann, C. R. Solis, J. A. Negrón-sotomayor et al., The maintenance of high genetic relatedness in multi-queen colonies of social wasps, Queen number and sociality in insects, pp.153-170, 1993.

R. Ihaka and R. Gentleman, R: a language for data analysis and graphics, J Comput Graph Stat, vol.5, pp.299-314, 1996.

K. Johnson, E. Duval, M. Kielt, and C. Hughes, Male mating strategies and the mating system of great-tailed grackles, Behav Ecol, vol.11, pp.132-141, 2000.

T. Kikuchi, R. Yoshioka, and N. Azuma, Effects of worker manipulation on the sex ratio of a Japanese ant species, Myrmecina nipponica, Ecol Res, vol.17, pp.717-720, 2002.

B. Manly, Randomization, bootstrap and Monte Carlo methods in biology, 1997.

N. J. Mehdiabadi, H. K. Reeve, and U. G. Mueller, Queens versus workers: sex ratio conflict in eusocial Hymenoptera, Trends Ecol Evol, vol.18, pp.88-93, 2003.

A. P. Møller and M. D. Jennions, Testing and adjusting for publication bias, Trends Ecol Evol, vol.16, pp.580-586, 2001.

A. P. Møller and M. D. Jennions, How much variance can be explained by ecologists and evolutionary biologists?, Oecologia, vol.132, pp.492-500, 2002.

U. G. Mueller, Haplodipoidy and the evolution of facultative sex ratios in a primitively eusocial bee, Science, vol.254, pp.442-444, 1991.

P. Nonacs, Ant reproductive strategies and sex allocation theory, Q Rev Biol, vol.61, pp.1-21, 1986.

P. Nonacs, Sex ratio determination within colonies of ants, Evolution, vol.40, pp.199-204, 1986.

K. M. Noonan, Sex-ratio of parental investment in colonies of social wasp Polistes fuscatus, Science, vol.199, pp.1354-1356, 1978.

P. Packer and R. E. Owen, Relatedness and sex ratio in a primitively eusocial halictine bee, Behav Ecol Sociobiol, vol.34, pp.1-10, 1994.

P. Pamilo, Sex allocation and queen-worker conflict in polygynous ants, Behav Ecol Sociobiol, vol.27, pp.31-36, 1990.

P. Pamilo and R. Rosengren, Sex ratio strategies in Formica ants, Oikos, vol.40, pp.24-35, 1983.

P. Pamilo and P. Seppä, Reproductive competition and conflicts in colonies of the ant Formica sanguinea, Anim Behav, vol.48, pp.1201-1206, 1994.

L. Passera, A. S. Vargo, E. L. Keller, and L. , Queen control of sex ratio in fire ants, Science, vol.293, pp.1308-1310, 2001.

R. J. Paxton, P. A. Thorén, A. Estoup, and J. Tengö, Queen-worker conflict over male production and the sex ratio in a facultatively polyandrous bumblebee, Bombus hypnorum: the consequences of nest usurpation, Mol Ecol, vol.10, pp.2489-2498, 2001.

M. Pearcy and A. S. , Local resource competition and sex ratio in the ant Cataglyphis cursor, Behav Ecol, vol.17, pp.569-574, 2006.

D. C. Queller and J. E. Strassmann, Kin selection and social insects, Bioscience, vol.48, pp.165-175, 1998.

D. C. Queller, J. E. Strassmann, C. R. Solis, C. R. Hughes, and D. M. Deloach, A selfish strategy of social insect workers that promotes social cohesion, Nature, vol.365, pp.639-641, 1993.

F. Ratnieks, K. R. Foster, and T. Wenseleers, Conflict resolution in insect societies, Annu Rev Entomol, vol.51, pp.581-608, 2006.

W. R. Rice and S. D. Gaines, Extending nondirectional heterogeneity tests to evaluate simply ordered alternative hypotheses, Proc Natl Acad Sci, vol.91, pp.225-226, 1994.
DOI : 10.1073/pnas.91.1.225

URL : http://www.pnas.org/content/91/1/225.full.pdf

Y. Roisin and A. S. , Split sex ratios in perennial social hymenoptera: a mixed evolutionary stable strategy from the queens' perspective?, Am Nat, vol.162, pp.624-637, 2003.

M. S. Rosenberg, D. C. Adams, and J. Gurevith, MetaWin: statistical software for meta-analysis, 2000.

R. Rosenthal, Meta-analytic procedures for social research, Thousand, 1991.

K. G. Ross and L. Keller, Ecology and evolution of social organization: insights from fire ants and other highly eusocial insects, Annu Rev Ecol Syst, vol.26, pp.631-656, 1995.

H. Rosset and M. Chapuisat, Sex allocation conflict in ants: when the queen rules, Curr Biol, vol.16, pp.328-331, 2006.

H. Rosset and M. Chapuisat, Alternative life-histories in a socially polymorphic ant, Evol Ecol, vol.21, pp.577-588, 2007.

G. Schino, Birth sex ratio and social rank: consistency and variability within and between primate groups, Behav Ecol, vol.15, pp.850-856, 2004.

B. C. Sheldon and S. A. West, Maternal dominance, maternal condition, and offspring sex ratio in ungulate mammals, Am Nat, vol.163, pp.40-54, 2004.

R. E. Stark, Sex-ratio and maternal investment in the multivoltine large carpenter bee Xylocopa sulcatipes, 1992.

, Ecol Entomol, vol.17, pp.160-166

J. E. Strassmann, Female-biased sex ratio in social insects lacking morphological castes, Evolution, vol.38, pp.256-266, 1984.

L. Sundström, M. Chapuisat, and L. Keller, Conditional manipulation of sex ratios by ant workers: a test of kin selection theory, Science, vol.274, pp.993-995, 1996.

L. Sundström and F. Ratnieks, Sex ratio conflicts, mating frequency, and queen fitness in the ant Formica truncorum, Behav Ecol, vol.9, pp.116-121, 1998.

R. L. Trivers and H. Hare, Haplodiploidy and the evolution of the social insects, Science, vol.191, pp.249-263, 1976.

E. L. Vargo, Sex investment ratios in monogyne and polygyne populations of the fire ant Solenopsis invicta, J Evol Biol, vol.9, pp.783-802, 1996.

L. Walin and P. Seppä, Resource allocation in the red ant Myrmica ruginodis-an interplay of genetics and ecology, J Evol Biol, vol.14, pp.694-707, 2001.

P. S. Ward, Genetic relatedness and colony organization in a species complex of Ponerine ants. II. Patterns of sex ratio investment, Behav Ecol Sociobiol, vol.12, pp.301-307, 1983.

S. A. West, A. S. Griffin, and A. Gardner, Evolutionary explanations for cooperation, Curr Biol, vol.17, pp.661-672, 2007.

S. A. West and B. C. Sheldon, Constraints in the evolution of sex ratio adjustment, Science, vol.295, pp.1685-1688, 2002.

S. A. West, D. M. Shuker, and B. C. Sheldon, Sex-ratio adjustment when relatives interact: a test of constraints on adaptation, Evolution, vol.59, pp.1211-1228, 2005.

D. Yanega, Caste determination and differential diapause within the first brood of Halictus rubicundus in New York (Hymenoptera, Halictidae), Behav Ecol Sociobiol, vol.24, pp.97-107, 1989.