Behavioral Ecology and Sociobiology

, Volume 70, Issue 1, pp 133–143 | Cite as

The relative importance of spatial proximity, kin selection and potential ‘greenbeard’ signals on provisioning behaviour among helpers in a cooperative bird

  • Paul G. McDonald
  • Lee Ann Rollins
  • Stephanie Godfrey
Original Article


Many hypotheses have been proposed to account for cooperative behaviour, with those favouring kin selection receiving the greatest support to date. However, the importance of relatedness becomes less clear in complex societies where interactions can involve both kin and non-kin. To help clarify this, we examined the relative effect of indirect versus key direct benefit hypotheses in shaping cooperative decisions. We assessed the relative importance of likely reciprocal aid (as measured by spatial proximity between participants), kin selection (using molecular-based relatedness indices) and putative signals of relatedness (vocal similarity) on helper/helper cooperative provisioning dynamics in bell miners (Manorina melanophrys), a species living in large, complex societies. Using network analysis, we quantified the extent of shared provisioning (helping at the same nests) among individual helpers (excluding breeding pairs) over three seasons and 4290 provisioning visits, and compared these with the location of individuals within a colony and networks built using either genetic molecular relatedness or call similarity indices. Significant levels of clustering were observed in networks; individuals within a cluster were more closely related to each other than other colony members, and cluster membership was stable across years. The probability of a miner helping at another’s nest was not simply a product of spatial proximity and thus the potential for reciprocal aid. Networks constructed using helping data were significantly correlated to those built using molecular data in 5 of 10 comparisons, compared to 8 of 10 comparisons for networks constructed using call similarity. This suggests an important role of kinship in shaping helping dynamics in a complex cooperative society, apparently determined via an acoustic ‘greenbeard’ signal in this system.


Cooperative breeding Direct benefits Inclusive fitness Indirect benefits Population viscosity Social networks analysis 



Nick and Joan Hoogenraad and the La Trobe University Wildlife Reserve kindly allowed fieldwork on their land. Maria Pacheco, Luc te Marvelde and Jonathon Wright assisted with fieldwork. Simon Griffith provided facilities for molecular analyses, with Anna Kopps and Serena Lam assisting with laboratory work. We thank two anonymous reviewers, Prof. Theo Bakker and Dr. Bernhard Voelkl for providing helpful comments on earlier drafts. LAR was funded through an ARC Discovery Project grant to Simon Griffith at Macquarie University (DP1094295), SG through a Discovery Early Career Researcher Award from the Australian Research Council (DE120101470) and PM by the University of New England and a Biotechnology and Biological Sciences Research Council grant to Jonathan Wright at Bangor University (5/S19268).

Compliance with ethical standards

Ethical approval

All applicable international, national and/or institutional guidelines for the care and use of animals were followed. All procedures performed in studies involving animals were in accordance with the ethical standards of the institution or practice at which the studies were conducted. This article does not contain any studies with human participants performed by any of the authors. Research was approved by the La Trobe University Animal Ethics committee (AEC01/19(L)/V2), the Department of Sustainability and Environment (licence 10002082) and the Australian Bird and Bat Banding Scheme (A2259), who also provided leg bands.


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Copyright information

© Springer-Verlag Berlin Heidelberg 2015

Authors and Affiliations

  • Paul G. McDonald
    • 1
    • 2
  • Lee Ann Rollins
    • 3
    • 4
    • 5
  • Stephanie Godfrey
    • 6
  1. 1.Centre for Behavioural and Physiological Ecology, ZoologyUniversity of New EnglandArmidaleAustralia
  2. 2.School of Biological SciencesUniversity of WalesBangorUK
  3. 3.Deakin UniversityGeelongAustralia
  4. 4.School of Life and Environmental SciencesCentre for Integrative EcologyGeelongAustralia
  5. 5.Department of Biological SciencesMacquarie UniversitySydneyAustralia
  6. 6.School of Veterinary and Life SciencesMurdoch UniversityPerthAustralia

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