Evolution of the sterile caste.

Biased sex ratios are expected to affect the conditions of the evolution of worker behaviour in malehaploid populations. In a subsocial, malehaploid species an association to female-biased brood sex ratios favours the evolution of worker behaviour. There are various reasons to expect such an association. The critical worker efficiency threshold required for worker behaviour to evolve is lower when looked at from the viewpoint of the mother than from that of the daughter. In this mother-daughter conflict, the mother can expect some help from her other offspring to resolve the conflict in her favour. In semisocial species the sex ratio biases have a less significant role and the worker behaviour can evolve more easily in malehaploid than in diploid populations. Furthermore, limited dispersal (due to the risks attached to it) is expected to favour worker behaviour in malehaploid but inbreeding in diploid populations. In addition to the genetic mechanisms that determine the critical worker efficiency threshold, it is essential to pay attention to those ecological factors that affect the actual worker efficiency in nature. Several such factors exist but it is not easy to associate them with the common occurrence of eusocial species in the Hymenoptera.

[1]  E. Charnov Kin selection and helpers at the nest: Effects of paternity and biparental care , 1981, Animal Behaviour.

[2]  J. Seger Partial bivoltinism may cause alternating sex-ratio biases that favour eusociality , 1983, Nature.

[3]  F. Ratnieks,et al.  Importance of the Sting in the Evolution of Sociality in the Hymenoptera , 1989 .

[4]  E. Charnov,et al.  Some conceptual issues in the origin of eusociality , 1986, Heredity.

[5]  Non-linear benefits and the evolution of eusociality in the hymenoptera , 1981 .

[6]  R. Snelling 5 – Systematics of Social Hymenoptera , 1981 .

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

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

[9]  C. Starr,et al.  Enabling Mechanisms in the Origin of Sociality in the Hymenoptera—the Sting's the Thing , 1985 .

[10]  R. D. Alexander,et al.  The evolution of social behavior , 1974 .

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

[12]  T. Myles,et al.  Termite Eusocial Evolution: A Re-Examination of Bartz's Hypothesis and Assumptions , 1988, The Quarterly Review of Biology.

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

[14]  D. Queller The evolution of eusociality: Reproductive head starts of workers. , 1989, Proceedings of the National Academy of Sciences of the United States of America.

[15]  Y. Iwasa Role of sex ratio in the evolution of eusociality in haplodiploid social insects , 1981 .

[16]  M. Andersson,et al.  The Evolution of Eusociality , 1984 .

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

[18]  B. Charlesworth,et al.  Some models of the evolution of altruistic behaviour between siblings. , 1978, Journal of theoretical biology.

[19]  P D Taylor,et al.  Inclusive fitness models with two sexes. , 1988, Theoretical population biology.

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

[21]  G. C. Eickwort 3 – Presocial Insects , 1981 .

[22]  R. Craig,et al.  Sex ratio changes and the evolution of eusociality in the hymenoptera: simulation and games theory studies. , 1980, Journal of Theoretical Biology.

[23]  K. Aoki,et al.  One- and two-locus models of the origin of worker behavior in hymenoptera , 1981 .

[24]  Charles D. Michener,et al.  Comparative Social Behavior of Bees , 1969 .

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

[26]  W. Hamilton The genetical evolution of social behaviour. I. , 1964, Journal of theoretical biology.

[27]  Peter D. Taylor,et al.  Evolutionary stability in one-parameter models under weak selection , 1989 .

[28]  R. Trivers,et al.  Haploidploidy and the evolution of the social insect. , 1976, Science.

[29]  C. Michener,et al.  Were workers of eusocial hymenoptera initially altruistic or oppressed? , 1974, Proceedings of the National Academy of Sciences of the United States of America.

[30]  R. Craig PARENTAL MANIPULATION, KIN SELECTION, AND THE EVOLUTION OF ALTRUISM , 1979, Evolution; international journal of organic evolution.

[31]  C. Michener,et al.  Evolution of Sociality in Insects , 1972, The Quarterly Review of Biology.

[32]  R. Crozier 6 – Genetics of Sociality , 1979 .

[33]  D. Yanega,et al.  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.

[34]  R. Crozier Evolutionary Genetics of the Hymenoptera , 1977 .

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