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• W2077063398 abstract "Female hairy-faced hover wasps forage for the young of a dominant breeder, but some forage more than others. New research shows that helpers decide how much to help by looking to the future. Female hairy-faced hover wasps forage for the young of a dominant breeder, but some forage more than others. New research shows that helpers decide how much to help by looking to the future. A meerkat spends five minutes digging a hole the length of its entire body. With its head, literally, in the sand, it is vulnerable to attack from predators. When it finally captures a big juicy grub, instead of enjoying the fruits of its labour, the first thing it does is look for a pup to feed. The pup will usually not be its own but the offspring of the dominant pair in its group. Thanks to the glamorous world of nature documentaries the seemingly altruistic behaviour of helpers in group-living animals is widely familiar. But what is really going on here? We know that natural selection is acting on helpers to maximise their reproductive success, so why doesn't our meerkat keep the grub for itself? Cooperatively breeding animals such as meerkats are excellent model systems for understanding the evolution of cooperative behaviour and are widely studied for this reason [1Clutton-Brock T.H. Breeding together: kin selection and mutualism in cooperative vertebrates.Science. 2002; 296: 69-72Crossref PubMed Scopus (669) Google Scholar, 2Cockburn A. Evolution of helping behavior in cooperatively breding birds.Annu. Rev. Ecol. System. 1998; 29: 141-177Crossref Scopus (614) Google Scholar]. However, there are inevitable limitations on research carried out on wild mammals and birds — many of these species are long-lived, and disperse over long distances, making it difficult to follow individuals over their lifetimes. Jeremy Field and colleagues [3Field J. Cronin A. Bridge C. Future fitness and helping in social queues.Nature. 2006; 441: 214-217Crossref PubMed Scopus (94) Google Scholar] have recently reported a study that manages to get around many of these problems, on the delightfully named hairy-faced hover wasp, Liostenogaster flavolineata Cameron. This wasp has a breeding system more like meerkats, with reproductively viable helpers, than their more familiar hymenopteran cousins the ants, where helpers are sterile. The fundamental question raised by cooperative breeding systems is how do helpers increase fitness by investing in the production of another individual's offspring? One possibility is that the offspring may be a sibling or cousin, and so genes for cooperative behaviour are passed-on by the production of non-descendent relatives [4Hamilton W.D. The genetical evolution of social behaviour, I & II.J. Theor. Biol. 1964; 7: 1-52Crossref PubMed Scopus (9548) Google Scholar]. Although there is some support for this explanation, it is not universally applicable: there are huge differences in the amount of effort helpers invest that do not correspond to differences in relatedness [5Griffin A.S. West S.A. Kin discrimination and the benefits of helping in cooperatively breeding vertebrates.Science. 2003; 302: 634-636Crossref PubMed Scopus (309) Google Scholar]. Alternatively, helpers may be keeping an eye on the future: helping today may maximise their success at breeding tomorrow. This can occur if helping increases group size or increases the number of helpers available in the future when they eventually inherit the dominant breeding position [6Kokko H. Johnstone R.A. Clutton-Brock T.H. The evolution of cooperative breeding through group augmentation.Proc. R. Soc. Lond. B. 2001; 268: 187-196Crossref PubMed Scopus (415) Google Scholar, 7Shreeves G. Field J. Group size and direct fitness in social queues.Am. Nat. 2002; 159: 81-95Crossref PubMed Scopus (68) Google Scholar]. In many cases it can be helpful to think of non-breeding helpers as being in a queue for dominant breeding status [8Kokko H. Johnstone R.A. Social queuing in animal societies: a dynamic model of reproductive skew.Proc. R. Soc. Lond. B. 1999; 266: 571-578Crossref Scopus (245) Google Scholar]. In a nest of hairy-faced hover wasps you would typically find one to ten related females forming an orderly queue for the chance to reproduce (Figure 1). A single dominant female, which is also the oldest, will lay the eggs and almost never leave the nest. It is the job of the other females, below her in the queue, to go out on foraging trips to collect food for the larvae. The more foragers there are, the more offspring are produced, so by increasing the reproductive success of the dominant the helpers are increasing their fitness by producing relatives. So far so good, but previous studies have identified a potential conflict of interest arising from this sort of breeding system: there will be a trade-off between investing in the reproduction of a dominant today and saving your energy in case you get the chance to breed as a dominant in the future [9Cant M.A. Field J. Helping effort and future fitness in cooperatve animal societies.Proc. R. Soc. Lond. B. 2001; 268: 1959-1964Crossref PubMed Scopus (130) Google Scholar]. The extent of this conflict is predicted to depend firstly, on the position an individual occupies in the social queue, and secondly, on the size of the prize at the end — breeding success depends on the size of the group. Do helpers save their energy for the chance to breed? There is very little evidence available to answer this question, mainly because of the difficulties in obtaining data on lifetime reproductive success. The study by Field et al.[3Field J. Cronin A. Bridge C. Future fitness and helping in social queues.Nature. 2006; 441: 214-217Crossref PubMed Scopus (94) Google Scholar] is important, not only because the authors were able to obtain accurate measures of helping investment, but also because they went one step further and experimentally manipulated both the chance of obtaining fitness in the future and the amount of fitness that was up-for-grabs in the future. This allowed them to test two key theoretical predictions: first, that individuals near to the top of the queue should work less hard than individuals near the bottom of the queue; and second, that individuals should work less hard in larger groups. In both cases, individuals should work less hard when they have more to lose. In their first experiment, Field et al.[3Field J. Cronin A. Bridge C. Future fitness and helping in social queues.Nature. 2006; 441: 214-217Crossref PubMed Scopus (94) Google Scholar] simply promoted their target wasp by removing the wasp that was ranked above her in the queue. They then compared the helping effort before and after the manipulation by simply measuring the amount of time spent out of the nest on foraging missions. As predicted, when the chance of inheriting the breeding position was increased, the target wasp slacked off on her foraging duties. In their second experiment, the authors manipulated the size of the group and, therefore, the potential workforce available to help them raise their own offspring when they inherited the breeding position. All wasps were removed except for numbers one and two in the queue from groups that initially contained three to five females. The authors then compared the change in investment made by rank two females before and after the manipulation. As predicted, the wasp in the larger groups invested less than the wasp in the same position in the queue of the reduced group. They controlled for offspring to helper ratio by also removing some larvae from the manipulated groups. To what extent do the findings of Field et al.[3Field J. Cronin A. Bridge C. Future fitness and helping in social queues.Nature. 2006; 441: 214-217Crossref PubMed Scopus (94) Google Scholar] apply more generally, for example to vertebrate species with similar breeding systems? A recent study [10MacColl A.D.H. Hatchwell B.J. Determinants of lifetime fitness in a cooperative breeder, the long-tailed tit Aegithalos caudatus.J. Anim. Ecol. 2004; 73: 1137-1148Crossref Scopus (59) Google Scholar] that measured lifetime reproductive success in a cooperatively breeding bird, the long-tailed tit, showed that helping contributed a small proportion to lifetime fitness in birds that eventually went on to become breeders. There is no evidence, however, that tits behave like wasps and trade-off fitness from helping for breeding in the future. The reason for this is actually very obvious, as the future is very uncertain for long tailed tits: mortality between breeding seasons is 55% [11McGowan A. Woodburn R.J.W. Hatchwell B.J. The effect of helping behaviour on the survival of juvenile and adult long-tailed tits (Aegithalos caudatus).J. Anim. Ecol. 2003; 72: 491-499Crossref Scopus (48) Google Scholar]. There will be significant variation between species in the extent to which individuals are able to gauge the expected future benefits of helping. The hairy-faced hover wasp can assess the two most important parameters that determine expected future fitness: their position in the social queue and group size. In many species it is not known how helpers assess their social position or if they are able to do so with accuracy. If there is not an orderly queue to reproduce it may pay to carry-on helping in an uncertain future." @default.
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• W2077063398 title "Social Evolution: Lazy Wasps Look to the Future" @default.
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