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The Coexistence Division of labour and risk taking in the dinosaur ant, Dinoponera quadriceps Article (Published Version) Asher, Claire L, Nascimento, Fabio S, Sumner, Seirian and Hughes, William O H (2013) Division of labour and risk taking in the dinosaur ant, Dinoponera quadriceps. Myrmecological News, 18. pp. 121-129. ISSN 1994-4136 This version is available from Sussex Research Online: http://sro.sussex.ac.uk/id/eprint/46186/ This document is made available in accordance with publisher policies and may differ from the published version or from the version of record. If you wish to cite this item you are advised to consult the publisher’s version. Please see the URL above for details on accessing the published version. Copyright and reuse: Sussex Research Online is a digital repository of the research output of the University. Copyright and all moral rights to the version of the paper presented here belong to the individual author(s) and/or other copyright owners. To the extent reasonable and practicable, the material made available in SRO has been checked for eligibility before being made available. Copies of full text items generally can be reproduced, displayed or performed and given to third parties in any format or medium for personal research or study, educational, or not-for-profit purposes without prior permission or charge, provided that the authors, title and full bibliographic details are credited, a hyperlink and/or URL is given for the original metadata page and the content is not changed in any way. http://sro.sussex.ac.uk Myrmecological News 18 121-129 Vienna, March 2013 Division of labour and risk taking in the dinosaur ant, Dinoponera quadriceps (Hyme- noptera: Formicidae) Claire L. ASHER, Fabio S. NASCIMENTO, Seirian SUMNER & William O.H. HUGHES Abstract The success of social insects can be largely attributed to division of labour. In contrast to most social insects, many species with simple societies contain workers which are capable of sexual reproduction. Headed by one or a few reproductive individuals, subordinate workers form a dominance hierarchy, queuing to attain the reproductive role. In these species task allocation may be influenced by individual choice based on future reproductive prospects. Individuals with a better chance of inheriting the colony may be less likely to take risks and high-ranking workers that spend a greater amount of time in proximity to the brood may be able to increase the ability to police egg-laying by cheating subordinates. We investigated division of labour and risk taking in relation to dominance rank in the queenless ponerine ant, Dinoponera quadriceps, a species with relatively simple societies. Using behavioural observations, we show that high-ranking wor- kers spend more time performing egg care, less time foraging and are less likely to defend the nest against attack. High- rankers also spent a greater amount of time guarding and inspecting eggs, behaviours which are likely to improve detec- tion of egg laying by cheating subordinates. We also show that high-ranking workers spend a greater amount of time idle, which may help increase lifespan by reducing energy expenditure. Our results suggest that both risk-taking and egg- care behaviours are related to future reproductive prospects in D. quadriceps. This highlights a mechanism by which effective division of labour could have been achieved during the early stages of eusocial evolution. Key words: Eusociality, Dinoponera quadriceps, social insect, dominance hierarchy, Ponerinae, polyethism. Myrmecol. News 18: 121-129 (online 1 March 2013) ISSN 1994-4136 (print), ISSN 1997-3500 (online) Received 17 October 2012; revision received 21 January 2013; accepted 7 February 2013 Subject Editor: Helge Schlüns Claire L. Asher (contact author), Institute of Integrative and Comparative Biology, University of Leeds, Leeds, LS2 9JT, UK; Institute of Zoology, Zoological Society of London, Regent's Park, London, England, NW1 4RY, UK. E-mail: [email protected] Fabio S. Nascimento, Departmento de Biologia, FFCLRP, Universidade de Sao Paulo, Av Bandeirantes 3900, BR- 14040901 Sao Paulo, Brazil. Seirian Sumner, Institute of Zoology, Zoological Society of London, Regent's Park, London, England, NW1 4RY, UK. William O.H. Hughes, Institute of Integrative and Comparative Biology, University of Leeds, Leeds, LS2 9JT, UK. Introduction Social insects represent one of the pinnacles of social evo- cies, the worker caste is further divided into individuals who lution, and the evolution of eusociality is considered to be specialise in certain tasks for at least part of their adult life one of the major transitions in evolution (MAYNARD SMITH (WILSON 1974, ROBINSON 1992, WILSON 2000). The majo- & SZATHMARY 1995). Understanding how such highly co- rity of social insect species lack morphologically differen- operative societies evolved has been a key area of interest tiated castes (WILSON 1974) and in these societies labour for evolutionary biologists since Darwin, who highlighted is often divided according to age, via temporal polyethism the apparent paradox of worker behaviour (DARWIN 1859). (WILSON 1974, TOFTS 1991, BESHERS & FEWELL 2001). One of the primary characteristics of eusocial societies is Most commonly, temporal polyethism involves younger division of labour (ROBINSON 1992, BESHERS & FEWELL workers performing tasks within the nest, and older workers 2001), a stable variation within a colony in the tasks that leaving the nest to forage (BESHERS & FEWELL 2001). This individuals perform (BESHERS & FEWELL 2001). Division results in individuals with a shorter life expectancy per- of labour is believed to have been a key factor in the suc- forming the more dangerous outdoor tasks (KAY & RIS- cess of social insects, increasing efficiency and maximis- SING 2005, MORON & al. 2008). ing resource use (WILSON 1974, WILSON 1985, PAGE & In contrast to species with large, complex societies, MITCHELL 1998). All social insect colonies show reproduc- species with small, relatively simple societies can some- tive division of labour, where a relatively small number of times contain workers who are physically capable of sex- individuals (queens) are responsible for reproduction, while ual reproduction, a state which is most likely ancestral in the other colony members (workers) rear the young, for- wasps and bees but which is secondarily derived in a num- age and maintain the nest (ROBINSON 1992). In some spe- ber of ant species (PEETERS 1991, FIELD & al. 2000). Re- production is still dominated by one or a few individuals, The dinosaur ant, Dinoponera quadriceps, has small, however, and younger subordinate workers queue to take simple, queenless societies, in which workers form a short, over the reproductive role (PEETERS 1991, SHREEVES & linear hierarchy behind the alpha (PEETERS & CREWE 1984, FIELD 2002, FIELD & CANT 2009). MONNIN & PEETERS 1999). When the alpha dies she is re- Queuing for reproduction results in differences in the placed by a high-ranked subordinate, most commonly the future reproductive prospects of different individuals in beta, and high-ranking workers themselves are frequently the colony (CANT & FIELD 2001, FIELD & CANT 2009). In replaced by newly emerged workers, resulting in age-based these societies individual workers may select tasks based hierarchy in which low-ranking workers are the oldest col- on the costs and benefits associated with them (CANT & ony members (MONNIN & PEETERS 1999). Observations FIELD 2001). Since high-ranking workers have a greater suggest that foraging and nest maintenance may tend to chance of future reproduction (PARDI 1948, MONNIN & be carried out by lower ranked individuals and brood care PEETERS 1998, 1999) we might expect them to be less like- by higher ranked individuals (MONNIN & PEETERS 1999), ly to engage in foraging and nest defence (CANT & FIELD in keeping with hierarchical position affecting division of 2001, FIELD & CANT 2009), tasks that are likely to have labour, but the data in support of this is still quite limited. a high mortality rate (VISSCHER & DUKAS 1997, FIELD & Here we carry out a detailed examination of the relationship CANT 2009). High-ranking individuals may also increase between dominance rank and behaviour in D. quadriceps. their chances of future reproduction by spending a greater Using observations of 24 behaviours and experimental sti- amount of time idle, or by increasing body condition, for mulation of nest defence we test the hypothesis that indi- example by increasing time spent grooming (FIELD & CANT viduals exhibit different behaviours in relation to their cur- 2009). However, to our knowledge these hypotheses have rent and future reproductive prospects. The effects of age not yet been thoroughly tested in a species in which wor- and rank are confounded in this species, therefore we do ker reproduction is secondarily derived. not attempt to identify the mechanism but merely whether Although subordinate workers in species with simple behaviour is related to reproductive potential. Specifically, societies may be physically capable of reproduction, there we predict that (1) high-ranking workers show lower en- is in fact little opportunity for them to gain direct repro- ergy expenditure and avoid dangerous tasks such as for- ductive fitness because in some species (e.g., ants), males aging and nest defence, thereby increasing their chances of show no interest in mating with subordinates (PEETERS future reproduction, and (2) high-rankers spend a greater 1991). However, unmated
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