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University of Groningen Sociable Schedules Helm, Barbara View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by University of Groningen University of Groningen Sociable schedules Helm, Barbara; Piersma, Theunis; Van der Jeugd, Henk Published in: Animal Behavior DOI: 10.1016/j.anbehav.2005.12.007 IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from it. Please check the document version below. Document Version Publisher's PDF, also known as Version of record Publication date: 2006 Link to publication in University of Groningen/UMCG research database Citation for published version (APA): Helm, B., Piersma, T., & Van der Jeugd, H. (2006). Sociable schedules: interplay between avian seasonal and social behaviour. Animal Behavior, 72(2), 245-262. https://doi.org/10.1016/j.anbehav.2005.12.007 Copyright Other than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons). Take-down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons the number of authors shown on this cover page is limited to 10 maximum. Download date: 12-11-2019 ANIMAL BEHAVIOUR, 2006, 72, 245e262 doi:10.1016/j.anbehav.2005.12.007 REVIEWS Sociable schedules: interplay between avian seasonal and social behaviour BARBARA HELM*,THEUNISPIERSMA†‡ & HENK VAN DER JEUGD†§ *Max Planck Institute for Ornithology yAnimal Ecology Group, Centre for Ecological and Evolutionary Studies zDepartment of Ecology and Evolution, Royal Netherlands Institute for Sea Research (NIOZ) xSOVON Dutch Centre for Field Ornithology (Received 26 June 2004; initial acceptance 15 September 2004; final acceptance 6 December 2005; published online 21 June 2006; MS. number: RV-51R) Timing is essential in seasonally changing habitats. Survival and reproduction are enhanced through precise adjustment to environmental conditions. Avian seasonal behaviour, that is, diverse activities associated with reproduction, moult and migration, has an endogenous basis and is ultimately linked to changes in environmental factors such as food supply. However, behaviour occurs in social contexts, and interactions with conspecifics are intimately linked to seasonal activities. Time programmes set the stage for social behaviour, which in turn fine-tunes seasonal activities. We propose that avian schedules are genuinely ‘sociable’: birds communicate seasonal behaviour by both intentional and inadvertent infor- mation transfer and negotiate it in competitive and cooperative interactions. Studying the interplay be- tween seasonal and social behaviour can add to our understanding of animal behaviour, including mechanisms by which birds could cope with changing environmental conditions. Ó 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved. Birds can undertake spectacular seasonal journeys; for major, and blue tits, Cyanistes caeruleus, time their clutches example, Arctic terns, Sterna paradisaea, almost circle the precisely to capitalize on the narrow peak in the availabil- globe each year (Alerstam 1990; Berthold 2001). Migra- ity of their primary nestling food, caterpillars (e.g. Perrins tions can be successful only if they coincide precisely 1970; Visser et al. 1998). Even slight mismatches between with favourable conditions, not only at target locations breeding and food peak can impose measurable costs: but also at staging areas en route. Mistimed migration im- spring weather changes have unsettled the phase relation poses high survival and fitness costs, such as when birds between predictive daylength information and food peak, encounter severe weather conditions (e.g. James 1956; thus imposing energetic and fitness costs on tits (Visser Brown & Brown 2000) or find reduced food resources at et al. 1998; Thomas et al. 2001). Although less extensively staging sites (Moore et al. 1995; Baker et al. 2003). The studied, other annual events, notably moult, often also oc- temporal precision of migratory birds has encouraged cur at precise times (e.g. Newton 1966; Stresemann & Stre- studies of their timing skills for almost a millennium semann 1967; Hahn et al. 1992; Jenni & Winkler 1994; (Gwinner & Helm 2003). Siikama¨ki et al. 1994; Nilsson & Svensson 1996). The need for accurate timing is not confined to Seasonal activities take place in social contexts, and migrants, however. Residents, such as great tits, Parus social interactions are a crucial aspect of annual cycles. Avian social behaviour, defined as including all within- species interactions, rarely remains constant over time. Correspondence: B. Helm, Max Planck Institute for Ornithology, Von- Many species are solitary for part of the year, pair up for der-Tann-Str. 7, D-82346 Andechs, Germany (email: [email protected]. breeding and flock together for migration and wintering de). T. Piersma is at the Royal Netherlands Institute for Sea Research (NIOZ) and Animal Ecology Group, Centre for Ecological and Evolu- (Rappole 1995; Greenberg & Salewski 2005). Such transi- tionary Studies, University of Groningen, PO Box 14, NL-9750 AA tions can be instantaneous: for example migratory species Haren, The Netherlands. H. van der Jeugd is at the SOVON Dutch Cen- often claim territories and try to attract mates immediately tre for Field Ornithology, Rijksstraatweg 178, 6573 DG Beek-Ubbergen, upon arriving at the breeding grounds. Avian species The Netherlands. differ widely in the plasticity and timing of their social 245 0003e3472/06/$30.00/0 Ó 2006 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved. 246 ANIMAL BEHAVIOUR, 72,2 behaviour. Reproductive ‘partnerships’ range from perma- (Gwinner 1986; Piersma 2002b). However, in nature, birds nent mate attendance to breeding partners that reunite for usually synchronize with the external year in reference to only a few hours every year (Black 1996a, b). Similarly, ter- timing cues. Rowan (1926) showed the importance of ritorial behaviour differs in timing and extent between photoperiod, the annual change in daylength for seasonal species. Common stonechats, Saxicola torquatus, tolerate behaviour. Daylength has now been identified to provide conspecifics during migration, but otherwise defend pair important ‘initial predictive information’ for timing of territories throughout the year (Ko¨nig et al. 2002; Urqu- numerous avian species (Hahn et al. 1992, 1997; Hau hart 2002); more sociable species are territorial for only et al. 1998; Wingfield & Jacobs 1999). Within the ‘win- short periods of the year (e.g. red knots, Calidris canutus: dows’ set by endogenous programmes, schedules are influ- Whitfield & Brade 1991). Highly social species, such as enced by other proximate factors (Gwinner 1999). Many waterfowl, vigorously defend nest sites and feeding areas species adjust timing in response to temperature, weather within a colony during the breeding season but may and food supply (e.g. Lack 1943, 1950, 1968; Perrins 1970; favour kin with which they associate in complex social Drent & Daan 1980; Bairlein & Gwinner 1994; Maney interactions (Andersson & A˚ hlund 2000; van der Jeugd et al. 1999; Piersma 2002a; Leitner et al. 2003). Seasonal et al. 2002). The physiological basis for behavioural behaviour is also affected by the availability of commodi- change is often endogenously programmed, as evidenced ties, such as nestboxes, which prime reproductive func- by annual fluctuations in seasonal activities and related tions (Gwinner et al. 2002; Wingfield & Silverin 2002). key hormones (Gwinner 1986; Wingfield & Marler 1988). Comparatively little is known about the effects of social For example, aggression and plasma testosterone of young interactions, however, with the possible exception of African stonechats held in Germany rose spontaneously sexual cycles (e.g. Lewis & Orcutt 1971; Gwinner 1986, when free-living conspecifics established territories (Ko¨nig 1999; Wingfield & Marler 1988; Hahn et al. 1997; Goldmann et al. 2002). et al. 2004). In this paper we aim to integrate research on timing and The way in which birds respond to temporal informa- social interactions. We first discuss timing mechanisms tion is influenced by their circannual programmes, which and social interactions from behavioural and evolutionary function as ‘periodically changing dispositions to respond perspectives. We propose that avian schedules are genu- to environmental cues’ (Gwinner 1999, page 2367). The inely ‘sociable’: seasonal activities are inadvertently and current phase of a bird’s circannual clock determines actively communicated and are often negotiated in con- how temporal information, for example daylength, is flict and cooperation. We illustrate the interplay between interpreted. Thus, short days generally accelerate seasonal seasonal and social behaviour by selected examples activities in the autumn but delay schedules in spring (e.g. throughout the annual cycle. A more detailed yet still Farner & Gwinner 1980; Hahn et al. 1992, 1997). In Fig. 1 incomplete review of literature is given in Table 1.We we propose how environmental factors, including social conclude by offering perspectives on studying sociable
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