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Habitat Selection of the Cooperative Breeding Taiwan Yuhina (Yuhina

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Habitat Selection of the Cooperative Breeding Taiwan Yuhina (Yuhina Zoological Studies 44(4): 497-504 (2005) Habitat Selection of the Cooperative Breeding Taiwan Yuhina (Yuhina brunneiceps) in a Fragmented Forest Habitat Pei-Fen Lee1, Sheng-Feng Shen1,2, Tzung-Su Ding3, Chyi-Rong Chiou3, and Hsiao-Wei Yuan3,* 1Institute of Ecology and Evolutionary Biology, and Department of Life Science, National Taiwan University, Taipei, Taiwan 106, R.O.C. E-mail: [email protected] 2Present address: Department of Neurobiology and Behavior, Cornell University, Ithaca, NY 14853, USA E-mail: [email protected] 3School of Forestry and Resource Conservation, National Taiwan University, Taipei, Taiwan 106, R.O.C. E-mail: [email protected] (TSD), [email protected] (CRC), and [email protected] (HWY) (Accepted July 22, 2005) Pei-Fen Lee, Sheng-Feng Shen, Tzung-Su Ding, Chyi-Rong Chiou, and Hsiao-Wei Yuan (2005) Habitat selection of the cooperative breeding Taiwan Yuhina (Yuhina brunneiceps) in a fragmented forest habitat. Zoological Studies 44(4): 497-504. We used multilevel analyses (individual habitat selection and population structure) to study edge effects on the Taiwan Yuhina (Yuhina brunneiceps), an endemic subtropical species at Meifeng, central Taiwan. The 95% kernel home range area was used to study habitat preferences, and the source-sink threshold was calculated to determine whether this yuhina population is self-sustainable. Within the study site, woods, open spaces, orchards, buildings, and pond habitats were preferred in that order. Within the home ranges, orchard habitats were preferred to open spaces. Larger breeding groups had larger home ranges with more trees and open spaces. Nest sites were closer to forest edges than random points were. The density of Taiwan cherry trees was significantly higher within home ranges than available areas, especially , near edges, suggesting the fruit s potential value to breeding yuhinas. Although the yuhinas suffered high rates of nest failure at Meifeng, this population still seemed to be self-sustaining due to the long breeding season with multiple broods and a cooperative breeding strategy. http://zoolstud.sinica.edu.tw/Journals/44.4/497.pdf Key words: Edge, Fragmentation, Home range, Source-sink population dynamics. Habitat fragmentation and habitat loss are and Villard 2001, Flaspohler et al. 2001), guild- considered major threats to biodiversity (Wilson specific responses (e.g., Stouffer and Bierregaard 1992). One unavoidable consequence of habitat 1995), and, theoretically, life history-specific fragmentation is an increase in edge habitats, responses (e.g., Donovan and Thompson 2001) to which in turn affects abiotic and biotic processes in edge habitats. How individuals perceive patchi- the landscape (Murcia 1995). Nesting birds may ness and how they move among fragments influ- disproportionately favor the edges of forest patch- ence how they are affected by fragmentation es due to the availability of forest cover and abun- (Wiens 1994, Ims 1995, Andreassen et al. 1998). dant food, but may suffer high nest predation as a Therefore, understanding the pattern of habitat result. Therefore, edges may become“ecological selection, together with the behavioral basis for traps”leading to severe population declines in the selection, can help explain variations in edge edge-nesting species (Gates and Gysel 1978). effects. Few studies have combined individual- The effects of habitat fragmentation are not level mechanisms with a population-level demo- homogeneous. There are species-specific graphic structure. Even fewer, if any, multilevel responses (e.g., Warkentin et al. 1995, Bourque studies have been conducted in tropical and sub- *To whom correspondence and reprint requests should be addressed. Tel: 886-2-23660235. E-mail: [email protected] 497 498 Zoological Studies 44(4): 497-504 (2005) tropical areas, where ecological contexts (e.g., ecology. By 2001, 97% of the adults in our study higher predation risks and longer breeding sea- site were color-banded. We closely monitored the sons) and life histories (e.g., multiple broods) may breeding success of 6, 11, and 13 groups in 1998, widely differ from those of temperate species. 2000, and 2001, respectively. Each morning of the These differences may have profound influences 2001 breeding season (early Mar. to late Aug.), we , on a species responses to edge habitats (Martin walked along a fixed route that covered the entire 1996, Russell 2000, Stutchbury and Morton 2001). study site, weather permitting. When we encoun- The Taiwan Yuhina (Yuhina brunneiceps), tered a banded group, we followed the group and , with monomorphic sexes, is a passerine bird plotted each member s location on a fine-scale endemic to Taiwan (Cibois et al. 2002). It is widely map every 5 min until contact was lost. To facili- distributed in forests from 800 to 3000 m in eleva- tate mapping, we established a grid system with tion (Koh and Lee 2003). Yuhinas are the only reference points every 20 m at the study site. known passerine species to consistently adopt a In 2001, 91 nests were found from 20 breed- joint-nesting strategy in a large fraction of nests ing groups for nesting analyses. We searched for (Vehrencamp and Quinn 2004). Breeding groups yuhina nests in both natural and man-made habi- include 2-7 individuals composed primarily of non- tats and mapped the location of each nest. We kin. Therefore juvenile delayed-dispersal is not were confident that most of the nests within the observed (Yuan et al. 2004). Living in groups typi- study site had been discovered, as yuhinas are an cally grants each individual increased protection active and conspicuous species, and every breed- from predators and increased chances of finding ing group was closely followed. food (Krebs and Davies 1993). In White-throated Magpie-Jays (Calocitta formosa), a cooperatively breeding species, it has been shown that larger groups possess larger territories containing more food, which contributes to higher reproductive suc- cess (Langen and Vehrencamp 1998). In this paper, we examined 1) the influence of fragmentation on yuhinas, at the individual level, with respect to (a) habitat selection and (b) the behavioral basis of habitat selection in terms of nest site and food resources; 2) this yuhina popu- , lation s possible status as a source population (in terms of meta-population dynamics (Pulliam 1988); and 3) variations in home range qualities among breeding groups of different sizes. MATERIALS AND METHODS Behavioral observations We conducted this study at Meifeng Highlands Experiment Farm of National Taiwan University in Nantou County, central Taiwan (24 05'N, 121 10'E, elevation 2150 m; Fig. 1). The° study site,° established for horticultural research purposes in 1961, a 50 ha area mixed with green- houses, temperate-fruit orchards, meadows, 100 50 0 100 M Japanese Cryptomeria (Cryptomeria japonica) N Ponds Orchards Open spacas plantations, and various ornamental trees sur- W E Buildings Woods rounded by natural forests composed mainly of S Fagaceae and Lauraceae. Fig. 1. Study area and land cover map at Meifeng, the Since 1995, we have banded adult and juve- Highlands Experiment Farm of National Taiwan Univ., Nantou nile yuhinas at Meifeng to study their breeding County, central Taiwan. Lee et al. -- Habitat Selection by Taiwan Yuhina 499 Although the diet of yuhinas varies with the by the density of the 3 major food plants within the seasons, it consisted primarily of fruits, nectar, and home range. Those plants shared by home insects within the study site. During the breeding ranges of multiple breeding groups were equally season, yuhinas mainly foraged on 3 plant divided by the number of groups and assigned to species: nectar (Jan.-Apr.) and fruits (May-June) of each home range. Taiwan cherry (Prunus campanulata), fruits (May- Two levels of use-availability design (Johnson July) of Eastern debregeasia (Debregeasia orien- 1980), i.e., utilization vs. home range and home talis), and nectar (Aug.-Sept.) of mistletoe (Taxillus range vs. available area, were conducted to lonicerifolius) (Liu 1999). We mapped the loca- assess habitat selection by yuhinas. We deter- tions of these 3 species within the study site and mined the available area for each group separately used their abundances as food resource indexes by generating 100 m circular buffers around all , for the yuhinas. group members occurrence locations by ArcGIS (McClean et al. 1998, Jones 2001). Because pro- Habitat data analysis portions of habitat types were not independent, we used compositional analyses to determine the A digitized habitat map of the study site was habitat preferences of yuhinas (Aebischer et al. created by interpreting an aerial photo (1:5000, 0.5 1993). x 0.5 m in spatial resolution, taken in 2000 by the Agricultural and Forestry Aerial Survey Institute of Demographic data analysis the Taiwan Forestry Bureau, Council of Agriculture, Executive Yuan) using ArcGIS 8.1 (ESRI 2001). The source-sink threshold, as the number of Five major habitat types were identified by land juveniles per pair per year, was calculated accord- cover and land use: woods, orchards, buildings, ing to the following equation (Trine 1998) for 1998, open spaces, and ponds (Fig. 1). Wooded habitat 2000, and 2001: was defined as forest patches and hedgerow strips 2＊ adult mortality with trees more than 3 m in height and included Source-sink threshold = 1-juvenile mortality primary forests and Cryptomeria plantations. Orchard habitats were plantations of temperate The adult mortality rate was estimated by the fruit trees, mostly apples, peaches, and pears. overwintering return rate. Because juveniles com- Building habitat included greenhouses, residences, monly disappear from Meifeng (78%, Yuan et al. and offices. Open space habitat included scat- 2004) and we have observed banded juveniles tered open spaces with shrubs < 3 m in height, which had dispersed as far as 7 km away, direct scattered individual trees, grass, bare ground, estimation of the juvenile mortality rate was trails, concrete or asphalt roads, and nearby areas.
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