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Egg Rejection Behavior and Clutch Characteristics of the European Greenfinch Introduced to New Zealand

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Egg Rejection Behavior and Clutch Characteristics of the European Greenfinch Introduced to New Zealand Chinese Birds 2012, 3(4):330–338 ORIGINAL ARTICLE DOI 10.5122/cbirds.2012.0037 Egg rejection behavior and clutch characteristics of the European Greenfinch introduced to New Zealand Peter SAMAŠ 1,, Lenka POLAČIKOVÁ 1,2, Mark E. HAUBER 3, Phillip CASSEY 4, Tomáš GRIM 1 1 Department of Zoology and Laboratory of Ornithology, Palacký University, 17. listopadu 50, CZ–771 46 Olomouc, Czech Republic 2 Department of Pathology and Parasitology, Faculty of Veterinary Medicine, University of Veterinary and Pharmaceutical Sciences, Palackého 1-3, CZ–612 42 Brno, Czech Republic 3 Department of Psychology, Hunter College and the Graduate Center of the City University of New York, 695 Park Avenue, New York, NY 10065, USA 4 School of Earth and Environmental Sciences, University of Adelaide, SA 5005, Australia Abstract Animal populations, with a known history of introduction events, provide opportunities to study the dynamics of how rapid shifts in ecological context affect behavioral (e.g., responses to brood parasitism) and life-history (e.g., clutch and egg parameters) traits. We studied the European Greenfinch (Carduelis chloris) introduced to New Zealand, regarding foreign-egg rejection behaviors and also compared their clutch characteristics with data from the source populations in the United Kingdom. Although previously this species had been considered an unsuitable host for the Common Cuckoo (Cuculus canorus), and not impacted by selection pressure associated with brood parasit- ism, we found that Greenfinches in our study population were able to eject experimental eggs at low frequencies. In contrast, nest desertion rates were similar in experimentally parasitized and control unmanipulated nests, implying that nest desertion is not an antiparasite adaptation in this species. Contrary to previous studies, we did not find significant differences in clutch and egg sizes between introduced and source populations. This study emphasizes (1) the importance of using control treat- ments in studies of host responses to experimental parasitism, (2) including apparently unsuitable hosts of brood parasites, and (3) meta-replicating prior studies to further the process of gaining and validating scientific knowledge. Keywords antiparasite behavior, avian brood parasitism, Carduelis chloris, clutch size, meta-replication, nest desertion Introduction more, 2011). Obligate brood parasitic birds, including the Common Cuckoo (Cuculus canorus; hereafter: Avian brood parasitism provides a powerful study sys- Cuckoo), and their hosts provide ideal models to study tem with challenging questions regarding the direction strong reciprocal selection pressures, because parasit- and speed of coevolutionary arms races, including host ism imposes moderate-to-severe costs on host parents selection, antagonistic interspecies interactions, and that provision unrelated young in their nests and hosts perceptual mimicry (Davies, 2011; Kilner and Lang- evolve antiparasitic traits (Davies, 2011). Such coevo- lutionary arms races have resulted in a diverse set of Received 30 November 2012; accepted 10 December 2012 quantifiable differences in the behaviors of both typi- cal and potential host populations that are either in Author for correspondence (Peter Samaš) contact, or in isolation, with a brood parasite (Davies E-mail: [email protected] and Brooke, 1989; Lahti, 2005; Hale and Briskie, 2007; www.chinesebirds.net Peter Samaš et al. Egg rejection in European Greenfinch 331 Stokke et al., 2007; Grim et al., 2011). genetic changes and may significantly influence clutch Research effort devoted to brood parasitism in birds characteristics, behavioral and sensory traits and other has increased dramatically in recent decades (Grim, aspects of introduced populations (Briskie and MacK- 2007). However, most studies have focused only on suit- intosh, 2004; Congdon and Briskie, 2010). In contrast, able Cuckoo hosts (Davies, 2011), whereas antiparasitic several studies have detected divergent changes in the behaviors of rare (Moksnes and Røskaft, 1992; Honza breeding characteristics of the Greenfinch between the et al., 2004; Procházka and Honza, 2004) or unsuitable source population in the UK and the introduced popu- (Davies and Brooke, 1989; Moksnes et al., 1991; Grim lation in NZ, including lower clutch size (Niethammer, et al., 2011) hosts have been tested only occasionally. 1970; MacMillan, 1985; Evans et al., 2005) and smaller Studying apparently unsuitable Cuckoo hosts provides egg size (Cassey et al., 2005) in NZ. These changes important insights for our understanding of host- support general observations that passerines in the parasite coevolution. For example, variation in the be- Northern Hemisphere have higher clutch sizes, fewer havioral responses of unsuitable hosts to experimental breeding attempts per season and/or a shorter breeding parasitism can simulate conditions at the start of brood season in contrast to the Southern Hemisphere regions parasitism in a novel (i.e., so far unused) host, and also (Martin, 1996; Samaš et al., 2013). give information on the phylogenetic distribution of Nevertheless, methodologically independent rep- behavioral traits that can serve as pre-adaptations for lication of whole studies (so called “meta-replication”) is defenses against brood parasites (Moskát et al., 2003a). critically important for the validity and generality of Species which have some allopatric populations, and scientific information (Johnson, 2002; Kelly, 2006). other sympatric populations, with brood parasites, pro- Therefore, additionally to testing Greenfinch responses vide valuable opportunities to study potential species- to experimental brood parasitism in allopatry with the level flexibility of antiparasitic strategies (Davies and Cuckoo, we also studied critical parameters of Green- Brooke, 1989; Stokke et al., 2007; Grim et al., 2011). Im- finch breeding biology in NZ (clutch and egg sizes). portantly, almost all studies on the effects of sympatry Traits of reproductive investment, including clutch size vs. allopatry with a brood parasite on host responses and egg size, are important components of avian para- were conducted in regions for which the length of sym- site-host arms races because clutch and egg sizes may patry or allopatry was unknown, i.e., reliable historical be related to individual host selection by female para- information on how long parasites were absent or co- sites and egg rejection decisions by hosts, and clutch habiting with hosts is missing in most cases (Davies and size may evolve divergently between host populations Brooke, 1989; Røskaft et al., 2002; Grim et al., 2011). sympatric vs. allopatric with parasites (e.g., Soler et al., Rare exceptions include the work of Lahti (2005) who 2001; Hauber, 2003; Servedio and Hauber, 2006; White knew accurately the length of allopatry in populations et al., 2007). Increase of clutch size might also act as a of Village Weaverbirds (Ploceus cucullatus) introduced defensive trait against brood parasites. For example, from Africa to the islands of Mauritius and Hispaniola, Svensson and Råberg (2010) and Soler et al. (2011) sug- and Hale and Briskie (2007) who knew the exact length gested that individual hosts better tolerate interspecific of allopatry in populations of several European avian parasitism by laying larger clutches. species introduced to New Zealand (see also Samaš et The Greenfinch is considered an unsuitable host for al., 2011). the Cuckoo because it feeds its nestlings mostly with Here, we take advantage of the known length of com- plant seeds (Davies and Brooke, 1989; Moksnes et al., plete allopatry with Cuckoos in the European Green- 1991). However, expectations on host suitability based finch (Carduelis chloris; hereafter: Greenfinch); this spe- on nestling diet composition have been shown empiri- cies was introduced from the United Kingdom (hereaf- cally to be misleading in certain cases (Martín-Gálvez ter: UK) to New Zealand (hereafter: NZ) between 1863 et al., 2005; Grim, 2006). Indeed, there are several doc- –1868 (Thomson, 1922). Although less than 100 indi- umented reports of Cuckoo chicks being successfully viduals were introduced, today the Greenfinch is com- raised by the Greenfinch (Seel and Davis, 1981). Clearly, mon throughout NZ and there is no genetic evidence more studies are needed to understand Greenfinch- for severe inbreeding or genetic drift in this species Cuckoo interactions in the context of brood parasitism, (Merilä et al., 1996; see also Briskie and MacKintosh, for example, to test whether flexible foraging and diet 2004). This is important as bottlenecks could lead to selection by Greenfinches, the flexibility of young Cuck- © 2012 Beijing Forestry University and China Ornithological Society 332 Chinese Birds 2012, 3(4):330–338 oos digestion, or the interaction of both these factors 1989), and show smaller clutch sizes and/or smaller egg can explain variation in the Cuckoo chick’s potential sizes in NZ in the southern hemisphere than in north- survival in Greenfinch nests (see also Grim et al., 2011). ern UK source populations (Cassey et al., 2005). No study has specifically tested the behavior of Greenfinch regarding egg rejection behavior in NZ Methods (cf. Boulton and Cassey, 2006; Hale and Briskie, 2007). In addition, only two European studies have reported All breeding and experimental data were collected dur- responses of the Greenfinch to experimental brood ing October and November 2007–2008 in the city of parasitism. Davies and Brooke (1989)
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