Nest-Site Microhabitat Association of Red-Billed Leiothrix in Subtropical
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Journal of Natural History ISSN: 0022-2933 (Print) 1464-5262 (Online) Journal homepage: http://www.tandfonline.com/loi/tnah20 Nest-site microhabitat association of red-billed leiothrix in subtropical fragmented forest in central China: evidence for a reverse edge effect on nest predation risk? Zhiqiang Zhang, Donghan Hou, Yuan Xun, Xuewen Zuo, Daode Yang & Zhengwang Zhang To cite this article: Zhiqiang Zhang, Donghan Hou, Yuan Xun, Xuewen Zuo, Daode Yang & Zhengwang Zhang (2016): Nest-site microhabitat association of red-billed leiothrix in subtropical fragmented forest in central China: evidence for a reverse edge effect on nest predation risk?, Journal of Natural History, DOI: 10.1080/00222933.2015.1130869 To link to this article: http://dx.doi.org/10.1080/00222933.2015.1130869 Published online: 17 Feb 2016. Submit your article to this journal View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tnah20 Download by: [University of Montana] Date: 19 February 2016, At: 22:24 JOURNAL OF NATURAL HISTORY, 2016 http://dx.doi.org/10.1080/00222933.2015.1130869 Nest-site microhabitat association of red-billed leiothrix in subtropical fragmented forest in central China: evidence for a reverse edge effect on nest predation risk? Zhiqiang Zhanga,b, Donghan Houb, Yuan Xunc, Xuewen Zuoc, Daode Yangb and Zhengwang Zhanga aMinistry of Education Key Laboratory for Biodiversity Science and Ecological Engineering, College of Life Sciences, Beijing Normal University, Beijing, China; bCollege of Forestry, Central South University of Forestry and Technology, Changsha, China; cConservation Institute of Daweishan Nature Reserve, Forestry Bureau of Liuyang City, Liuyang, China ABSTRACT ARTICLE HISTORY Previous studies of nest-site selection on a fine scale may reveal Received 2 March 2015 limiting resources within habitat types. The red-billed leiothrix Accepted 20 November 2015 (Leiothrix lutea Scopoli, 1786) is a common bird species that lives KEYWORDS in the subtropical forests of Asia. Despite many reports of this Microhabitat; nest-site species from introduced populations, little information has been selection; edge effects; obtained from its native range. From 2011 to 2013, we studied red-billed leiothrix; China nest-site selection of red-billed leiothrix at micro-scales in Daweishan Nature Reserve, Hunan Province, central China. A total of 363 nests were found in five vegetation types. We mea- sured the habitat variables and constructed nest-site selection models for nests found in the forest and scrub-grassland. Among the 18 variables measured in the forest, six variables were selected to construct the nest-site selection model: distance to forest edge (DTE), distance to water (DTW), vegetation comprehensive cover- age, tree coverage, bamboo coverage and shrub height. According to Akaike’s information criterion, the best model con- sisted of five of these variables (excluding vegetation comprehen- sive coverage), and distance to forest edge, distance to water, tree coverage and bamboo coverage had negative effects on nest-site selection. In scrub-grassland, the DTE, DTW, and bush coverage (BUC) were selected from the 13 variables measured, and, accord- ingly, the best model consisted of DTE and BUC. Model averaging suggested that BUC had a positive effect on nest-site selection. In Downloaded by [University of Montana] at 22:24 19 February 2016 contrast, DTE has a reverse effect. In addition, DTE differed sig- nificantly between successful and failed nests in forest and scrub- grassland. More successful nests were found near the forest edge. Taken together, these findings emphasise the power of fine-scale habitat selection models in identifying relevant habitat variables with a significant effect on preferred habitat and eventually, breeding success. CONTACT Zhengwang Zhang [email protected] Ministry of Education Key Laboratory for Biodiversity Science and Ecological Engineering, College of Life Sciences, Beijing Normal University, Beijing, 100875, China. © 2016 Taylor & Francis 2 Z. ZHANG ET AL. Introduction Birds’ preferred habitat associations are expected to maximise their fitness by increasing the reproduction of individuals via nest-site selection (Martin 1988). Previous studies have suggested that selection patterns are often based on the synthesis of multiple factors and cues (Kristan et al. 2007), such as the availability of food, water and mates; predators; climate; and vegetation structure including vegetation height, density and cover (Martin 1993a; van Gils et al. 2006; Tellería and Pérez-Tris 2007; Patthey et al. 2012; Li et al. 2015). Furthermore, habitat characteristics preferred by birds also occur simulta- neously on several specific scales (Manly et al. 2002). Different specific scales have their habitat characteristics (Apps et al. 2001), and thus, birds may shift their preferences among different scales for particular activities (Leopold and Hess 2013). For example, some passerines usually select microhabitats for nest establishment (Latif et al. 2011; Murray and Best 2014). The microhabitat at a finer scale may reveal the limiting resource with the real needs of the topographic features or vegetation characteristics in the breeding period (Apps et al. 2001). The red-billed leiothrix Leiothrix lutea (Scopoli, 1786) is a small babbler (Timallidae) that inhabits dense bush in evergreen broadleaf and pine forests (Collar and Robson 2007). Its native distribution range is in Asia, including Southern China, North East Pakistan, North India, Nepal, Bhutan, North Myanmar and North Vietnam (Collar and Robson 2007; Zheng 2011). Historically, this species was highly appreciated and kept in captivity universally due to its colourful plumage and melodious songs (Cheng 1963). Thus, it has been introduced to Hawaii (Fisher and Baldwin 1947), Japan (Eguchi and Masuda 1994), the Island of Réunion and a number of scattered localities in Europe including France, Italy, Germany and Spain (Herrando et al. 2010; Farina et al. 2013), where the life-history traits of the introduced populations have been studied and reported partially during the past 20 years (Eguchi and Masuda 1994; Ralph et al. 1998; Amano and Eguchi 2002a, 2002b; Eguchi and Amano 2008; Herrando et al. 2010; Farina et al. 2013; Tojo and Nakamura 2014; Yang et al. 2014). However, limited information has been obtained in the native range of Asia, except Ma et al. ( 2010) and Zhou et al. (2012b), who described the nesting habitats of this babbler briefly in China, and suggested that red-billed leiothrix nested in dwarf bamboo or short bushes shaded by trees or shrubbery on slopes or nearby farmland. Recently, compared with the habitat-selection theory and methodology in early Downloaded by [University of Montana] at 22:24 19 February 2016 animal studies (MacArthur and Pianka 1966; Verner et al. 1986), many researchers have been paying more attention to employing statistical models in Akaike’s information criterion (AIC) framework and the cross-validation criterion (CVC) framework to demon- strate habitat association patterns and reveal nest-site selection preferences (Horne and Garton 2006; Wang et al. 2012; Murray and Best 2014). Thus, more ecologists have been identifying the heterogeneity of environmental factors by comparing used habitat with unused habitat or available habitat (Jones 2001; Crampton and Sedinger 2011; Schmidt et al. 2014). Moreover, nesting near the habitat edge, as an obvious habitat characteristic of nest- sites preferred by the red-billed leiothrix, has been described in most previous studies (Amano and Eguchi 2002a; Herrando et al. 2010; Ma et al. 2010). However, birds nesting along the forest edges usually faced higher nest predation than those nesting in the JOURNAL OF NATURAL HISTORY 3 forest core area, which is known as the so-called edge effect (Gates and Gysel 1978; Batáry and Báldi 2004; Vetter et al. 2013). Many studies have shown that the edge effect was extremely obvious on avian breeding success in fragmented habitat in temperate forests (Söderström 1999; Batáry and Báldi 2004). However, an inverse edge effect on avian nest predation has been found in tropical fragmented forest on the basis of some experimental studies (Carlson and Hartman 2001; Spanhove et al. 2009; Sedláček et al. 2014). Some recent studies have indicated that several species of passerines also prefer to nest near the forest edge in the subtropical fragmented forest in Asia, such as Emei Shan liocichla Liocichla omeiensis (Riley, 1926;Fu2011), Chinese babax Babax lanceolatus (Verreaux, 1870; Xu et al. 2012), and yellow-throated bunting Emberiza elegans (Temminck and Laugier, 1835; Chen et al. 2015). Unfortunately, only a few studies have suggested an extremely obvious edge effect on avian nest predation in the subtropical forest. Thus, more research is needed to understand the habitat association and breeding success of subtropical forest birds, such as red-billed leiothrix in its native range (Amano and Eguchi 2002a; Herrando et al. 2010). From 2011 to 2013, we examined the nest-site selection and nest success of red-billed leiothrix in a fragmented forest at Daweishan Nature Reserve (DSNR) in central China, which is one of the important native ranges for this species. We expected to identify the nest-site microhabitat association pattern and cues preferred by the red-billed leiothrix and to determine whether the red-billed leiothrix preferred to nest near the habitat edge, and the presence or absence of an edge effect on nest predation risk. Materials and methods Study site Field work was conducted at DSNR, a provincial nature reserve of 6681 ha in the northern section of the Luoxiao Mountains, which is located in the northeast of Liuyang County, Hunan Province, central China (28°20′54″–28°28′ 47″N, 114°01′51″– 114°12′52″E; Figure 1). DSNR has an east-to-west orientation consistent with the tectonic line, with an elevation disparity from 230 to 1608 m. It is in the humid monsoon climate zone in the central subtropical region (with an annual average temperature of 13°C, and annual rainfall of 1800–2000 mm). It has a short rainy and humid spring (Chen et al.