Threat-Sensitive Anti-Predator Defence in Precocial Wader, the Northern Lapwing Vanellus Vanellus
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acta ethol DOI 10.1007/s10211-016-0236-1 ORIGINAL PAPER Threat-sensitive anti-predator defence in precocial wader, the northern lapwing Vanellus vanellus Natalia Królikowska1 & Jakub Szymkowiak1 & Rebecca Anne Laidlaw2,3 & Lechosław Kuczyński1 Received: 14 January 2016 /Revised: 26 April 2016 /Accepted: 6 May 2016 # The Author(s) 2016. This article is published with open access at Springerlink.com Abstract Birds exhibit various forms of anti-predator behav- rook throughout the breeding season, while defence against iours to avoid reproductive failure, with mobbing—observa- hooded crow, harriers and black-headed gull did not exhibit tion, approach and usually harassment of a predator—being clear temporal patterns. Mobbing behaviour of lapwings ap- one of the most commonly observed. Here, we investigate parently constitutes a flexible anti-predator strategy. The anti- patterns of temporal variation in the mobbing response exhib- predator response depends on predator species, which may ited by a precocial species, the northern lapwing (Vanellus suggest that lapwings distinguish between predator types vanellus). We test whether brood age and self-reliance, or and match mobbing response to the perceived hazard at dif- the perceived risk posed by various predators, affect mobbing ferent stages of the breeding cycle. We conclude that a single response of lapwings. We quantified aggressive interactions species may exhibit various patterns of temporal variation in between lapwings and their natural avian predators and used anti-predator defence, which may correspond with various generalized additive models to test how timing and predator hypotheses derived from parental investment theory. species identity are related to the mobbing response of lap- wings. Lapwings diversified mobbing response within the Keywords Harmtooffspringhypothesis .Parentalinvestment breeding season and depending on predator species. Raven theory . Predator recognition . Reproductive value of offspring Corvus corax, hooded crow Corvus cornix and harriers hypothesis evoked the strongest response, while common buzzard Buteo buteo, white stork Ciconia ciconia,black-headedgull Chroicocephalus ridibundus and rook Corvus frugilegus were Introduction less frequently attacked. Lapwings increased their mobbing response against raven, common buzzard, white stork and Predation constitutes the most important cause of reproductive failure in the majority of bird species (Ricklefs 1969; Montgomerie and Weatherhead 1988; Lima and Dill 1990; Electronic supplementary material The online version of this article (doi:10.1007/s10211-016-0236-1) contains supplementary material, Martin 1995). Birds engage in various forms of anti-predator which is available to authorized users. behaviour (Caro 2005) to enhance the chance of successful breeding. The most frequent are nest concealment and cryptic * Natalia Królikowska behaviours (Lima and Dill 1990), reducing the number of [email protected] eggs under increased risk of nest predation (Eggers et al. 2006), nesting near protector species (Lima and Dill 1990), 1 Department of Avian Biology and Ecology, Faculty of Biology, feigning injury behaviours (Caro 2005), changing their risk Adam Mickiewicz University, Umultowska 89, sensitivity depending on the risk and the option for re- 61-614 Poznań,Poland nesting (Schneider and Griesser 2015) and active nest de- 2 School of Biological Sciences, University of East Anglia, Norwich fence, like mobbing (Caro 2005; Lima and Dill 1990). Research Park, Norwich NR4 7TJ, UK Mobbing—observation, approach and usually harassment of 3 South Iceland Research Centre, University of Iceland, Bankavegi, a predator by prey birds with alarm calls and direct physical IS-800 Selfoss, Iceland attacks (Caro 2005)—may effectively deter predators from acta ethol nesting areas, thus ensuring higher offspring survival (Knight relatively scarce (Brunton 1990;Larsenetal.1996; Sordahl and Temple 1986;Caro2005; Kontiainen et al. 2009). 2004; Mathot et al. 2009). Precocial species allocate their However, mobbing also involves some costs, as it potentially parental investments differently and face different trade-offs increases predation risk to defending parents or even attracts than altricial ones due to differences in the parental care sys- other predators into an area due to its conspicuousness (Krama tem. Such differences may shape patterns of anti-predator re- and Krams 2005; Krams et al. 2007). Furthermore, mobbing is sponse. Moreover, the hypotheses derived from the parental costly in terms of time and energy expenditure, as it reduces investment theory were mostly considered separately (Brown the time available for other activities (Lima and Dill 1990; et al. 2006; Rhoades and Blumstein 2007;Mathotetal.2009; Caro 2005). Consequently, parents should trade off the invest- Crawford et al. 2012), and these studies explored patterns of ment in current reproduction (anti-predator behaviour) and the temporal variation in defence concerning a narrow range of ability to invest in the future (Clutton-Brock 1991; Stearns predator taxa. Evaluating brood defence behaviour throughout 1992). the breeding season in precocial species, having mobile and The relationship between investment in current and future dispersed chicks, is critical to gaining a wider understanding reproduction constitutes a life-history trade-off (Coleman and of anti-predator behaviour. Gross 1991; Stearns 1992; Ghalambor and Martin 2001). The majority of waders (including the northern lapwing Parental investment theory has yielded three hypotheses to Vanellus vanellus) consistently lay four eggs (Maclean 1972; explain patterns of temporal variation in eggs or chicks de- Walters 1984). Thus, in this group, there is no relationship fence during the breeding season. The ‘reproductive value of between defence intensity and clutch size or clutch volume offspring’ hypothesis assumes that the intensity of anti- (Rytkönen et al. 1995; Ruusila and Pöysa 1998; Kis et al. predator defence matches the reproductive value of offspring. 2000), contrary to altricial species, in which anti-predator re- Thus, parents should take greater risks and increase mobbing sponse often constitutes a function of brood size (Jónsson and intensity with increasing clutch and chick age (Andersson Gunnarsson 2010) or parents adjust the number of eggs ac- et al. 1980; Montgomerie and Weatherhead 1988). cording to the risk of predation (Eggers et al. 2006;Zanetteet According to the predictions of the ‘harm to offspring’ hy- al. 2011). In waders, the reproductive value of offspring in- pothesis, defence intensity should peak when offspring are creases with its age, but the offspring vulnerability likely more vulnerable to the temporary suspension of parental care peaks around hatching (Galbraith 1988; Montgomerie and (Dale et al. 1996; Listøen et al. 2000; Fernández and Llambías Weatherhead 1988; Brunton 1990; Johansson and Blomqvist 2013). The third hypothesis takes into account that for birds 1996; Weggler 2009). Chicks of waders (solely precocial spe- nesting in a multi-predator environment, different predators cies) are relatively self-reliant and demand markedly less pa- constitute different threats (Montgomerie and Weatherhead rental attention than altricial young (Walters 1982, 1984; 1988; Lima and Dill 1990; Palestis 2005;Lima2009). Some Schekkerman et al. 2009). The self-feeding activity of chicks predators threaten the broods only temporarily (i.e., they feed may expose them to predation (Schekkerman et al. 2009), and mainly either on eggs or chicks), while others may potentially parents likely defend them more vigorously than eggs, also depredate them at all stages of the breeding cycle (e.g. due to the nests of many species being relatively well Teunissen et al. 2008). The ‘threat-sensitive predator avoid- concealed (Šálek and Cepáková 2006). Older offspring that ance’ hypothesis suggests that individuals distinguish be- thermoregulate independently are more mobile and able to tween different types of risk (some predators are hazardous disperse (Weggler 2009), and thus have better survival and only for the offspring while other pose risk only for adults or reproductive prospects than younger ones. for both) and adjust anti-predator response to the perceived The northern lapwing (hereafter referred to as lapwing) hazard posed by a given predator (Helfman 1989;Schneider is a ground-nesting wader that primarily inhabits extensive- and Griesser 2013). Such a flexible anti-predator behaviour ly managed farmland dominated by pastures and marshes allows individuals to reduce the costs associated with brood or wet meadows with short vegetation. Populations of lap- defence and is exhibited by several bird species (e.g. wing have been declining in most European countries in Montgomerie and Weatherhead 1988; Mathot et al. 2009; recent decades (Krebs et al. 1999; Donald et al. 2001; Strnad et al. 2012). Newton 2004). These declines have resulted primarily from The majority of avian studies on offspring defence have low reproductive performance due to the interactive effects focused on altricial (Krama et al. 2012;reviewedin of habitat alterations and predation (Vickery et al. 2001; Montgomerie and Weatherhead 1988;Caro2005) or semi- Evans 2004;Wilsonetal.2004; MacDonald and Bolton precocial species (Clode et al. 2000;WhittamandLeonard 2008;Teunissenetal.2008). Red fox Vulpes vulpes and 2000; Palestis 2005). While it has been suggested that the corvids likely cause the majority