1. Sociality and kin selection in insects 鄭先祐 靜宜大學 生態學系

2. kin selection in insects2 Contents 9.1 introduction 9.1 introduction 9.2 The origin and evolution of eusociality 9.2 The origin and evolution of eusociality 9.3 The evolution of a stable reproductive skew 9.3 The evolution of a stable reproductive skew 9.4 Sex ratio evolution and kin conflict in social insects 9.4 Sex ratio evolution and kin conflict in social insects 9.5 conclusion 9.5 conclusion

3. kin selection in insects3 9.1 Introduction Eusocial society: alongside cooperative care of the brood and an overlap of adult generations, exhibit a reproductive division of labour. Eusocial society: alongside cooperative care of the brood and an overlap of adult generations, exhibit a reproductive division of labour. –Some members are specialized for reproduction (queens or kings), whereas others devote themselves to foraging, nest construction, defence and brood-reaing. –Reproductive altruism

4. kin selection in insects4 Eusocial insects Most eusocial insects belong to two orders, the Hymenoptera ( 膜翅目 ) (ants, bees and wasps) and the Isoptera (termites) ( 白蟻 ). Most eusocial insects belong to two orders, the Hymenoptera ( 膜翅目 ) (ants, bees and wasps) and the Isoptera (termites) ( 白蟻 ). –All ants and termites are eusocial, but many bees and wasps are solitary or exhibit other grades of social organization. A number of aphids (Hemiptera) are arguably also eusocial, as are some beetles and thrips. A number of aphids (Hemiptera) are arguably also eusocial, as are some beetles and thrips. Table 9.1 The major groups of eusocial insects and their traits. Table 9.1 The major groups of eusocial insects and their traits.

5. kin selection in insects5 Hamilton ’ s rule The idea at the heart of the modern understanding of the evolution of altruism and sociality is Hamilton ’ s theory of kin selection. The idea at the heart of the modern understanding of the evolution of altruism and sociality is Hamilton ’ s theory of kin selection. Social actions are ones that increase or decrease the offspring production of conspecifics. Social actions are ones that increase or decrease the offspring production of conspecifics. Hamilton ’ s rule, the gene for altruism undergoes selection if the condition r1b - r2c >0 is satisfied. Hamilton ’ s rule, the gene for altruism undergoes selection if the condition r1b - r2c >0 is satisfied. –r1 is the altruist ’ s relatedness to the beneficiary ’ s offspring –r2 is the altruist ’ s relatedness to its own offspring.

6. kin selection in insects6 West-Eberhard (1975), 將 Hamilton ’ s rule 轉換為 if rb – c > 0, where r is the relatedness of the altruist to the beneficiary. West-Eberhard (1975), 將 Hamilton ’ s rule 轉換為 if rb – c > 0, where r is the relatedness of the altruist to the beneficiary. Table 9.2 Relatedness levels in a social insect colony. Table 9.2 Relatedness levels in a social insect colony.

7. kin selection in insects7 9.2 The origin and evolution of eusociality 9.2.1 The necessity of kin selection 9.2.1 The necessity of kin selection 9.2.2 Hymenopteran eusociality and the haplodiploidy( 單套雙套 ) hypothesis 9.2.2 Hymenopteran eusociality and the haplodiploidy( 單套雙套 ) hypothesis 9.2.3 The origin of eusociality in the termites 9.2.3 The origin of eusociality in the termites

8. kin selection in insects8 The necessity of kin seleciton Reciprocal altruism is not ‘ true ’ altruism. Reciprocal altruism is not ‘ true ’ altruism. Eusociality usually entails true altruism. Eusociality usually entails true altruism. Hamilton ’ s rule shows that both genetic factors (affecting relatedness) and ecological ones (affecting benefit and cost) must be important in promoting eusocial evolution. Hamilton ’ s rule shows that both genetic factors (affecting relatedness) and ecological ones (affecting benefit and cost) must be important in promoting eusocial evolution. Eusocial evolution : Mutualism hypothesis vs. parental manipulation hypothesis Eusocial evolution : Mutualism hypothesis vs. parental manipulation hypothesis

9. kin selection in insects9 Hymenopteran eusociality and the haplodiploidy hypothesis Eusociality has evolved independently many times among the insects. Eusociality has evolved independently many times among the insects. Wilson (1971) estimated that there have been 11 origins of eusociality in the Hymenoptera, and one in the termites. Wilson (1971) estimated that there have been 11 origins of eusociality in the Hymenoptera, and one in the termites. A phylogeny based on mitochondrial DNA sequences suggests that the aphid family Hormaphididae evolved a a soldier caste at least five times. A phylogeny based on mitochondrial DNA sequences suggests that the aphid family Hormaphididae evolved a a soldier caste at least five times.

10. kin selection in insects10 Relatedness levels in the Hymenoptera Males are haploid and develop from unfertilized eggs, but females are diploid and develop from fertilized eggs. (haplo- diploidy) Males are haploid and develop from unfertilized eggs, but females are diploid and develop from fertilized eggs. (haplo- diploidy) 如此，整體 的 relatedness 比較高。 如此，整體 的 relatedness 比較高。 Haplo-diploidy hypothesis Haplo-diploidy hypothesis

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12. kin selection in insects12 9.3 The evolution of a stable reproductive skew Skew theory suggest several factors of this type. Skew theory suggest several factors of this type. Fig. 9.2 factors affect the stable level of reproductive skew. Fig. 9.2 factors affect the stable level of reproductive skew. – 促進 reproductive skew 的因素： Group productivity, ecological constraints, relatedness – 抑制的因素： subordinate fighting ability

13. kin selection in insects13 Skew evolution in the polistine wasps Reeve and Nonacs (1992) Polistes fuscatus wasps, skew reproduction, 有多 個雌蜂， alpha, beta Reeve and Nonacs (1992) Polistes fuscatus wasps, skew reproduction, 有多 個雌蜂， alpha, beta 偷偷移走一些 eggs 偷偷移走一些 eggs Alpha queens showed no consistent response to egg removal, but beta ones became more aggressive on average. Alpha queens showed no consistent response to egg removal, but beta ones became more aggressive on average.

14. kin selection in insects14 Skew evolution in the leptothoracine ants Bourke and Heinze (1994), polygynous colonies live in extended uniform habitats such as pinewoods. Bourke and Heinze (1994), polygynous colonies live in extended uniform habitats such as pinewoods. –Cost of dispersal are therefore relatively low. –All queens lay eggs and live together peaceably. (skew is low) Monogynous colonies live in a rocky area. Monogynous colonies live in a rocky area. –Cost of dispersal is high. –Permanently wingless, only one queen.

15. kin selection in insects15 9.4 Sex ratio evolution and kin conflict in social insects Relatedness asymmetry = relateness to systers / relatedness to brothers Relatedness asymmetry = relateness to systers / relatedness to brothers –0.75/0.25 = 3:1 (Table 9.2) –Sex ratio = 3:1 in favour of females. Tests of sex ratio theory in social Hymenoptera Tests of sex ratio theory in social Hymenoptera Table 9.4 Tests of sex ratio theory within monogynous, non-parasitic ant species. Table 9.4 Tests of sex ratio theory within monogynous, non-parasitic ant species. –Population sex investment ratio 通常都是偏向雌性 (>0.5) ，唯一有個例外 (0.36)

16. kin selection in insects16 Some colonies at mother-daughter associations should produce female-biased broods. Some colonies at mother-daughter associations should produce female-biased broods. Sundstrom (1994), monogynous population of wood ant (Formica truncorum), using allozyme analysis. Sundstrom (1994), monogynous population of wood ant (Formica truncorum), using allozyme analysis. –Some colonies had a multiply mated queen, produced mostly males –Others had a singly mated queen produced mainly females.

17. kin selection in insects17 How workers manipulate sex allocation Sundstrom et al (1996) shown that all queens in a monogynous population of Formica exsecta contributed a similar fraction of haploid eggs to their colony ’ s egg pool. Sundstrom et al (1996) shown that all queens in a monogynous population of Formica exsecta contributed a similar fraction of haploid eggs to their colony ’ s egg pool. –However, workers in colonies headed by singly mated queens raised a female bias of adult sexuals, –Whereas workers under multiply mated queens raised a male bias.

18. kin selection in insects18 9.5 conclusion Two unsolved issues: 1. The adaptive significance of within-colony kin discrimination 2. The evolution of multiple mating.

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