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note
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The haplodiploid sex determination system leads to a threefold relatedness between workers and their sisters (r = 0.75) as compared with that between workers and their brothers (r = 0.25).
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Kin selection theory states that individuals can pass on copies of their genes not only by producing offspring but also by helping nondescendent kin, such as siblings, to reproduce. By preferentially favoring their closest kin, organisms can maximize the number of their gene copies transferred to the next generation and thereby their inclusive fitness [W. D. Hamilton, J. Theor. Biol. 7, (1964)].
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+, each with 2 alleles). When data from all these markers were combined, only 2% (less than one colony) of double matings may have passed undetected, either because one patriline was not included in the sample or because of identical paternal genotypes at the diagnostic loci. Thirty-four queens were singly mated, 19 were doubly mated, 2 were triply mated, and 1 had mated with four males. The colonies with a multiply mated queen were combined in the further analyses.
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10544251129
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note
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2 = 1.03, df = 1, P > 0.2516).
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25
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10544231525
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note
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In all, 1209 workers and 503 males were analyzed. All males were haploid, and in all except one colony their genotype was consistent with them being the sons of the queen. The exceptional colony was probably orphaned, because no workers were produced and males of three different genotypes were found (which is consistent with workers producing the males). Furthermore, genotypes of males and workers revealed that two colonies had more than one queen. These three colonies were excluded from all analyses.
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26
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10544234679
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note
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Colonies headed by one singly mated queen have a higher relatedness asymmetry (3:1) than do colonies headed by a doubly (2:1) or triply mated queen (1.67: 1).
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27
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10544240819
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2 = 73/127, P > 0.2). The observed mating swarm sex ratio was 65% females (numerical sex ratio) and 71% females (investment sex ratio based on dry weights).
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To determine primary sex ratios, approximately 50 eggs (a small fraction of the total number of eggs) were collected at the onset of egg laying, between late April and mid-May, when sexual brood is produced [K. Gösswald, Die Waldameise Vol. 1 (AULA-Verlag, Wiesbaden, Germany; 1989)]. The number of chromosomes (2n = 52) [E. Hauschiteck-Jungen, and H. Jungen, Insect Soc. 23, 513 (1976)] was assessed (11) (blind with respect to colony identity and queen mating frequency) in at least 5 but usually 10 cells per egg. The large number of chromosomes sometimes precluded counts of all chromosomes, so eggs in which fewer than 26 chromosomes were consistently found were classified as males, whereas those with more than 40 chromosomes were classified as females. Only nests with at least 20 successfully sexed eggs were included in the analyses, so that two nests in 1994 and three in 1995 were discarded.
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Die Waldameise Vol. 1
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Gösswald, K.1
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37
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0001056437
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To determine primary sex ratios, approximately 50 eggs (a small fraction of the total number of eggs) were collected at the onset of egg laying, between late April and mid-May, when sexual brood is produced [K. Gösswald, Die Waldameise Vol. 1 (AULA-Verlag, Wiesbaden, Germany; 1989)]. The number of chromosomes (2n = 52) [E. Hauschiteck-Jungen, and H. Jungen, Insect Soc. 23, 513 (1976)] was assessed (11) (blind with respect to colony identity and queen mating frequency) in at least 5 but usually 10 cells per egg. The large number of chromosomes sometimes precluded counts of all chromosomes, so eggs in which fewer than 26 chromosomes were consistently found were classified as males, whereas those with more than 40 chromosomes were classified as females. Only nests with at least 20 successfully sexed eggs were included in the analyses, so that two nests in 1994 and three in 1995 were discarded.
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Hauschiteck-Jungen, E.1
Jungen, H.2
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38
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10544231510
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note
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We made efforts to collect eggs immediately after the onset of egg laying in April/May, when only sexual brood is laid. However, worker-destined eggs are laid from mid-May onward, causing a slight overlap in worker and sexual brood, so a few worker-destined eggs may have been included in the primary sex ratio sample. Hence, the primary sex ratios obtained here may slightly overestimate the proportion of females among the eggs, and consequently provide a conservative test of differences in primary and secondary sex ratios in the singly mated class. The sample sizes differ from those given in Table 1, because three colonies that were scored for primary sex ratios only produced worker brood in 1994.
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39
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10544234243
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note
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We thank J. J. Boomsma, A. F. G. Bourke, W. D. Brown, E. A. Herre, M. Milinski, D. Nash, T. Reuter, F. L. W. Ratnieks, P. Sherman, and three anonymous referees for comments on the manuscript; C. Chang, R. Forsman, P. Gertsch, A.-M. Mehmeti, I. Schmidt, and H. Åberg for assistance: and Tvärminne Zoological Station in Finland for working facilities. The study was funded by the Academy of Finland; Svenska Kulturfonden; Société Vaudoise des Sciences Naturelles; Swiss National Science Foundation; and Fondation du 450e anniversaire, University of Lausanne.
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