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The Importance of Landscape Structure for Nest Defence in the Eurasian Treecreeper Certhia Familiaris

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The Importance of Landscape Structure for Nest Defence in the Eurasian Treecreeper Certhia Familiaris Ornis Fennica 84:145–154. 2007 The importance of landscape structure for nest defence in the Eurasian Treecreeper Certhia familiaris Ari Jäntti, Harri Hakkarainen, Markku Kuitunen*, Jukka Suhonen Jäntti, A., Department of Biological and Environmental Science, P. O. Box 35 (YAC), Survontie 9, FI-40014, University of Jyväskylä, Finland Hakkarainen, H., Section of Ecology, Department of Biology, University of Turku, FI- 20014 Turku, Finland Kuitunen, M., Department of Biological and Environmental Science, P.O. Box 35 (YAC), Survontie 9, FI-40014, University of Jyväskylä, Finland. [email protected] (*Cor- responding author) Suhonen, J., Department of Biological and Environmental Science, P. O. Box 35 (YAC), Survontie 9, FI-40014, University of Jyväskylä, Finland. Current address: Section of Ecology, Department of Biology, University of Turku, FI-20014 Turku, Finland Received 19 April 2006, revised 9 August 2007, accepted 20 August 2007 Forest loss and fragmentation induces harmful ecological effects especially for species preferring mature forests. The Eurasian Treecreeper, Certhia familiaris, is highly special- ised in foraging on large tree trunks and can only occasionally forage outside of mature fo- rests. We quantified nest defence behaviour of Treecreeper parents toward a stuffed model of Great Spotted Woodpecker Dendrocopos major in central Finland. We used a Geographical Information System (GIS) to measure the landscape structure within a 200 m radius around the nest. We found that females with more fledged offspring gave alarm calls from farther away from the predator model than did females with fewer fledged off- spring. The alarming distance of females was longer when the forest patch around the nest was larger. In males, however, alarming distance decreased with increasing home patch size. It seems that forest loss may influence parental nest defence behaviour, which is one of the fundamental life-history traits in birds. The association between habitat characteris- tics and nest defence behaviour of birds need more attention to understand how human modified habitats affect bird breeding success. 1. Introduction success and other fitness-related consequences are committed to follow the constraints set by patch Destruction of natural habitats results in habitat size, habitat within patches, variable food supply, loss, edge effects, and habitat isolation. The last and inter-specific interactions, and changes in veg- two elements have been termed habitat fragmenta- etation structure and climatic conditions close to tion (Andrén 1994, Fahrig 1997, Schmiegelow & the edge (Gates & Gysel 1978, Møller 1988 & Mönkkönen 2002), and may have several harmful 1991, Robinson et al. 1995, Burke & Nol 2000, biological and ecological effects (Andrén 1994). Zanette et al. 2000). Bird species often show vari- In fragmented environments, behaviour, breeding able responses to forest loss and fragmentation 146 ORNIS FENNICA Vol. 84, 2007 (Schmiegelow & Mönkkönen 2002), but in this benefit of nest defence is increased brood survival, study we concentrate on a bird species which is but nest defence also has costs like energy expen- strongly associated with old growth forests. The diture and a risk of injury or even death (Mont- reproductive performance of mature forest birds gomerie & Weatherhead 1988). It could also de- has been shown to decrease with an increasing crease the future reproductive life of the parents level of habitat fragmentation (Gates & Gysel (Wallin 1987). The intensity of nest defence may 1978, Robinson et al. 1995, Kurki et al. 2000). be affected by many factors such as clutch and Modern forestry usually diminishes the quality brood size, offspring age, parental age, laying of forested environments as a living habitat for fo- date, weather and predator type and food condi- rest-dwelling species. It especially causes the loss tions (Knight & Temple 1986, Winkler & Wilkin- of mature forest habitats, particularly in the boreal son 1988, Clutton-Brock 1991, Hakkarainen & forest ecosystems. As a result of this rational forest Korpimäki 1994). planning, forest blocks are fragmented and sepa- We examined the effects of landscape structure rated by large clear-cuts and dense sapling stands on nest-defence behaviour in the Eurasian Tree- (Mykrä et al. 2000). This environmental alteration creeper Certhia familiaris (hereafter “Treecreep- increases the harmful effects on species that are er”). The Treecreeper is an excellent study object dependent on continuous and homogeneous wood for forest fragmentation studies because it breeds and forest cover (Niemelä 1999, Kouki & Väänä- in old-growth spruce (Picea abies) and mixed nen 2000). These species may have restricted abil- spruce and pine (Pinus sylvestris) coniferous fo- ity to disperse between suitable habitat patches rests (Kuitunen 1987, Kuitunen & Helle 1988, (Hansson 1992), and the energy expenditure of Aho et al. 1999). Further, its breeding densities are food and mate searching may fail in strongly frag- three times higher in old-growth forests than in mented landscapes (e.g. Rolstad & Wegge 1989). managed forests (Virkkala et al. 1994, Haila et al. Also, increased predation pressure in fragmented 1989). Also, this species is known to be extremely forest landscapes is exhaustively documented (e.g. sensitive to forest fragmentation, because it is ab- Kuitunen & Helle 1988, Andrén 1992, Kurki & sent from clear-cuts and saplings (Kuitunen & Linden 1995, Huhta 1995, Huhta et al. 2003, Helle 1988). Treecreepers breed in specially de- 2004). A general interest in associations between signed nest boxes in managed forests (Kuitunen forest habitat loss and fragmentation and forest 1987). Breeding parents need large trees for forag- bird communities has led to the publication of sev- ing (Suhonen & Kuitunen 1991a, Aho et al. 1997 eral studies from all over the world (Haila et al. a, b, Jäntti et al. 2001) where they search for ar- 1993, Faaborg et al. 1995, Freemark et al. 1995, thropods for their nestlings (Kuitunen & Törmälä Greenberg et al. 1997, Boulinier et al. 2001). In 1983, Suhonen & Kuitunen 1991b). The Tree- several studies, it has also been shown that forest creeper is a monogamous passerine bird showing fragmentation due to timber production has delete- biparental offspring care (Kuitunen & Suhonen rious effects on forest birds (e. g. Askins et al. 1989, Kuitunen et al. 1996) and nest defence, es- 1987, Jokimäki & Huhta 1996, Schmiegelow et al. pecially during the first brood (Jäntti et al. 2003). 1997, Luck 2002). For species experiencing Potential nest predators for Treecreepers in our strong habitat fragmentation, both empirical data study area are the Great Spotted Woodpecker (Doncaster et al. 1996, Jansson & Angelstam Dendrocopos major, Least Weasel Mustela nivalis 1999) and theoretical models predict that there are and Stoat Mustela erminea (Kuitunen & Alek- critical thresholds of habitat proportions within a nonis 1992). landscape at which various ecological processes In this study we had two aims. First, we studied change abruptly (O’Neill et al. 1989, Dytham whether parent Treecreeper nest defence behav- 1995, With & Crist 1995, Bascompte & Solé iour was related to number of nestlings and laying 1996). date, since the number of nestlings is an important Nest defence is one of the most important life- life-history trait. Second, because habitat loss and history traits in the reproductive investment of landscape fragmentation has often been associated birds and it increases the probability of successful with poor habitat quality we aimed to study pos- breeding (Blancher & Robertson 1982). The main sible associations between forest loss and frag- Jäntti et al.: Tree creeper nest defence in fragmented forests 147 mentation and parental nest defence behaviour. No lasted for five minutes. Generally, the arrival time study so far has documented the effects of land- was different for the male and female, so the trial scape structure on the nest defence behaviour of was performed separately for each parent. Arrival birds. time was the time from setting up the predator model until the parent arrived and started to dis- play (Jäntti et al. 2003). 2. Material and methods Mobbing rate was the attack or fluttering be- haviour of the Treecreeper towards the predator 2.1. Study area and general methods model during the five minute period. The distances of parents from the predator were measured during This study was conducted in 60–100 year old co- the trial, and after the trial the mean, minimum, and niferous and mixed managed forests in the vicinity maximum distances from the predator model were of Konnevesi Research Station in central Finland recorded. The mean distance from the predator (62º37’ N, 26º20’ E) during the summer of 1991. was the mean value of the bird to predator dis- There were about 60 forests with two special nest tances recorded every fifteen seconds during the boxes for Treecreepers, one for the first clutch and trial. The minimum distance was defined as the one for the second clutch (Kuitunen 1987). We shortest recorded distance of the Treecreeper to the used mist nets to trap breeding females at the nest model during the trial. The maximum distance was near the end of egg incubation, and males were the longest observed distance of the parent from trapped when they were feeding their offspring. the model during the trial. We also recorded the The birds were individually marked both with alu- number of parental visits into the nest box (Jäntti et minium and coloured rings. The throat of each par- al. 2003). ent was coloured to distinguish the females from The call rate was the mean number of tjii warn- the males. The adult birds were sexed by morpho- ing calls per minute during the five minute period. logical criteria, e.g. bill length, and the presence of At each nest, we waited for about 20 minutes to a brood patch in females (Suhonen & Kuitunen make sure the parents were present or absent. Dur- 1991a, b, Kuitunen unpublished data). ing the trial, we also observed the level of quieting that resulted after the parents’alarm call to address whether the high-pitched calls are enough to si- 2.2.
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