Patterns, correlates, and paternity consequences of extraterritorial foray behavior in the field sparrow (Spizella pusilla): an automated telemetry approach
In many territorial species, occasional movements beyond territory boundaries (extraterritorial forays) have been documented in many species. While many explanations for the occurrence of extraterritorial forays have been proposed, a logical and proposed function of extraterritorial forays is to engage in extra-pair copulations with extra-pair mates outside of their territories. We used an automated radio-telemetry system (ARTS) to examine the patterns and correlates of foray behavior of male and female field sparrows (Spizella pusilla) and investigated if forays were associated with extra-pair paternity (EPP) and cuckoldry. We found that male and female field sparrows regularly engaged in extraterritorial forays. In males, age and time of foraying (day vs. night) were important factors explaining foray rates (foray/h); older (ASY) males forayed more than younger (SY) males and, while we detected many nocturnal forays, most forays occurred during the day. For females, fertility stage and age appeared to be important in explaining foray rates; older females forayed more during pre-fertile period than fertile and post-fertile periods. Unlike foray rates, the duration of forays (min) was not explained by any of the variables examined. Surprisingly, despite the large number of forays documented (>3500), greater foray rates or duration of forays were not associated with higher probability of EPY for males or females or with cuckoldry in males. Forays may play a role in prospecting and acquiring information about their social and ecological environment, which ultimately may help them to achieve greater reproductive success, but not necessarily in the form of EPP.
Despite many territorial species are known to conduct extraterritorial forays (movements beyond their territory), very little is known about this behavior. We used an automated radio-telemetry system (ARTS) to examine the patterns, correlates, and paternity consequences of extraterritorial foray behavior in male and female field sparrows (Spizella pusilla). We documented more than 3500 forays and found that both male and female field sparrows regularly engaged in extraterritorial forays; however, different factors explain their foray rates (age, time of foraying (day vs. night), and fertility stage) but not the duration of forays. Surprisingly, greater foray rates or duration of forays were not associated with higher probability of EPY in males or females or with cuckoldry in males. Rather than exclusively acquiring extra-pair matings, forays likely serve multiple purposes, such as prospecting and acquiring information about their social and ecological environment, which ultimately may help individuals achieve greater reproductive success.
KeywordsExtraterritorial foray Extra-pair paternity Extra-pair copulations Mating systems Reproductive performance Automated radio-telemetry systems (ARTS)
We thank Jeff Brawn, Andy Suarez, and two anonymous reviewers for their comments on previous drafts of the manuscript. We thank Devin Kerr, Eric Swenson, Jessica Burton, Brian Pappadopoli, Zara John, Sarah Tomke, Ben Neece, Victor Zhang, Jonathon Jackson, Jill Deppe, Kyle Van den Bosch, and Scott Chiavacci for their help in the field and laboratory. We also thank the Vermilion County Conservation District for permission to conduct research at Kennekuk Cove County Park.
Compliance with ethical standards
The National Science Foundation (DDIG-14070801), Animal Behavior Society Student Research Grant, American Ornithologist Union Research Award, Illinois Ornithological Society Grants, Department of Natural Resources and Environmental Sciences travel awards, University of Illinois at Urbana-Champaign student opportunity grants, and the Illinois Department of Natural Resources (W-154-R-6).
Conflict of interest
The authors declare that they have no conflict of interest.
All procedures performed were in accordance with the ethical standards of the University of Illinois’ Institutional Animal Care and Use Committee under the IACUC protocol #10127 and the USGS bird banding permit (23577).
Informed consent was obtained from all individual participants included in the study.
- Birkhead TR, Møller AP (1992) Sperm competition in birds. Academic Press, LondonGoogle Scholar
- Brown JL (1964) The evolution of diversity in avian territorial systems. Wilson Bull 76:160–169Google Scholar
- Brown JL (1969) Territorial behavior and population regulation in birds. Wilson Bull 81:293–329Google Scholar
- Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information-theoretic approach. Springer, New YorkGoogle Scholar
- Carey M, Burhans DE, Nelson DA (1994) Field sparrow (Spizella pusilla). In: Poole A,Gill F (eds) The birds of North America, No. 103. The Academy of Natural Sciences, Philadelphia, Pennsylvania and The American Ornithologists’ Union, Washington, DCGoogle Scholar
- Danchin E, Heg D, Doligez B (2001) Public information and breeding habitat selection. In: Clobert J, E. Danchin E, Dhondt AA, Nichols J (eds) Dispersal. Oxford University Press, Oxford, pp 243–258Google Scholar
- Giraldeu LA (1997) The ecology of information use. In: Krebs JR, Davis NB (eds) Behavioural ecology. Blackwell Scientific, Oxford, pp 42–68Google Scholar
- Kempenaers B, Dhondt AA (1993) Why do females engage in extra-pair copulations? A review of hypotheses and predictions. Belg J Zool 123:93–103Google Scholar
- Kleven O, Marthinsen G, Lifjeld JT (2006) Male extraterritorial forays, age and paternity in the socially monogamous reed bunting (Emberiza schoeniclus). J Ornithol 147:468–473Google Scholar
- Littell RC, Milliken GA, Stroup WW, Wolfinger RD, Schabenberger O (2006) SAS for mixed models, 2nd edn. SAS Institute Inc, Cary, NCGoogle Scholar
- Pyle P (1997) Identification guide to north American birds, part 1. Slate Creek Press, Bolinas, CAGoogle Scholar
- R Development Core Team (2008) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna http://www.R-project.org, Urbana, USAGoogle Scholar
- Schuett W, Tregenza T, Dall SRX (2010) Sexual selection and animal personality. Biol. Rev. 85:217— 246Google Scholar
- Sellaeg DE (2014) Inferring female extra-pair mating behaviour from observed patterns of extra-pair paternity with a process-based model. The University of Bergen, Bergen, Norway, MSc thesisGoogle Scholar
- Silverman BW (1986) Density estimation for statistics and data analysis. Chapman and Hall, Urbana, USAGoogle Scholar