Biology Reference
In-Depth Information
for dominance, in both insects and vertebrates, is reviewed by Field and Cant (2009).
The social insects are discussed in more detail in Chapter 14.
Kin discrimination occurs in some cooperative breeding vertebrates, such as long-
tailed tits and the Seychelles warbler, but not in others, such as meerkats and
kookaburras. Cornwallis et al . (2009) show that this variation can be explained by
variation in the relative benefit of kin discrimination. Specifically, kin discrimination is
more likely when helping provides a greater benefit, and less likely when relatedness is
higher within groups (i.e. when indiscriminate helping within group will be directed at
relatives, so there is less to gain from discrimination).
Another potential example of by-product benefits is provided by the wasp Polistes
dominulus , where genetic analyses of natural nests have showed that 15-35%
of  subordinates were unrelated to the dominant female. In this species, the
helping  behaviour of these unrelated subordinates appears to be explained by the
direct fitness benefit obtained from nest inheritance (Queller et al ., 2000; Leadbeater
et al ., 2011).
Clutton-Brock (2009c) discusses the lack of empirical support for the importance of
reciprocity. Boyd and Richerson (1988) show that there are theoretical problems for
reciprocity when interactions are between more than two individuals. Stevens and
Hauser (2004) discuss why psychological mechanisms may prevent reciprocity in
animals. Raihani et al . (2010) provide an example of punishment across the sexes in
cleaner fish. Jander and Herre (2010) provide another example of enforcement in a
between species mutualism - fig trees sanction fig wasps that provide poor pollination.
Frank (2003) provides an overview of enforcement.
TOPICS FOR DISCUSSION
1. Discuss the relative merits of long-term field studies and field experiments for evaluating
the costs and benefits of cooperation.
2. It is clear that behaviours such as punishment or sanctions can favour cooperation. However,
if they are costly to perform, how can natural selection favour such enforcement mechanisms?
3. Read Krams et al . (2008), Russell and Wright (2009) and Wheatcroft and Krams (2009)
and discuss whether the mobbing behaviour of pied flycatchers is reciprocal cooperation.
What experiments would you carry out to resolve this controversy?
4. Why are cooperation and sexual selection so rarely studied in the same species (Boomsma,
2007)?
5. Discuss where the five explanations for cooperation given in Nowak (2006) fit onto
Fig. 12.2 or into Table 12.2.
6. 'A whole generation of scholars swallowed the line that the Prisoners' dilemma embodies
the essence of the problem of human cooperation' (Binmore, 2007, p. 18). Discuss.
7. Do any non-human organisms have a 'sense of fairness'?
8. Discuss the potential medical applications of bacterial behavioural ecology (Andre  &
Godelle, 2005; Brown et al ., 2009).
9. Discuss whether studies of human behaviour when playing economic games can be
applied to help tackle the problem of global warming (Milinski, 2006).
10. MacLean and Gudelj (2006) show how the free rider problem arises with respect to the
metabolism of sugar by yeast. How is the problem solved?
11. Can the soybean-rhizobia interaction be thought of as reciprocity?
12. Does free-loading occur in the fruiting body behaviour of microbes (Strassmann et al .
2000; Velicer et al . 2000; Buttery et al . 2009)?
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