Aeropalynological Study of Yangmingshan National Park, Taiwan

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1 Taiwania, 50(2): 80-92, 2005 Morphological Characters of Bird Species in Taiwan Hau-Jie Shiu (1), Tzung-Su Ding (2), Jia-En Sheu (1), Ruey-Shing Lin (1,3), Chau-Nien Koh (1,4), (1, 5) and Pei-Fen Lee (Manuscript received 18 March, 2005; accepted 12 April, 2005) ABSTRACT: We documented body measurements of birds captured during banding research in Taiwan by Wild Bird Society Taiwan. A total of 223 bird species with measurements of body mass, length of body, head, tail, bill and tarsus, and maximum wing span were compiled for the database. Body mass, length of bill, and length of tarsus of 85 species that had sample sizes > 20 were reported. In addition, we also compared our data with that from Dunning (1993). Among the ten species eligible for further analyses, we found that eight species showed significant differences in body mass between our data and that of Dunning (1993). Our results together with our recent studies (Lee et al, 2004; Ding et al., 2005) showed that compiling banding research data contribute to predict ecological parameters of bird studies in community, ecosystem and landscape levels. KEY WORDS: Bill length, body mass, geographic variation, Taiwan, tarsus length. INTRODUCTION Avian body size (e.g., mass and length) is one of the most accessible factors relevant to physiological processes, life history strategies, behavioral and ecological functions (Peters, 1983; Gaston et al., 2001). The body sizes of birds are frequently used in physiological and ecological studies (Schluter and Repasky, 1991; Dunning, 1993). In addition, body size also plays an important role in predicting the relationships of abundance and density (Griffiths, 1992; Blackburn and Gaston, 1997; Fa and Purvis, 1997; Ding et al., 2005), as well as species richness (Finlay et al., 1996; Gregory, 1998), ecosystem characteristics (Holling, 1992; Cumming and Havlicek, 2002), and macroecology (Gregory, 1998; Gaston and Blackburn, 2000; Cassey and Blackburn, 2004). Avian body size has been shown to be one of the accurate and less variant measures in birds (Peters, 1983), and is often recorded as mass (weight) or length. Dunning (1993), however, suggested that body mass is more useful and desirable than body length for most physiological and ecological studies. Body mass, unfortunately, could only be measured while birds are alive, and may not be available in many situations, which also explains why bird body length are well documented but body mass are not. 1. Institute of Ecology and Evolutionary Biology, National Taiwan University, 1, Roosevelt Road, Sec. 4, Taipei 106, Taiwan. 2. School of Forestry and Resource Conservation, National Taiwan University, 1, Roosevelt Road, Sec. 4, Taipei 106, Taiwan. 3. Endemic Species Research Institute, Council of Agriculture, 1, Ming-Shen East Road, Jiji, Nantou 552, Taiwan. 4. Taiwan Forestry Research Institute, Council of Agriculture, 53, Nanhai Road, Taipei 100, Taiwan. 5. Corresponding author. Tel: ; Fax: ; leepf@ntu.edu.tw

2 June, 2005 Shiu et al.: Avian database in Taiwan 81 To our knowledge, a comprehensive report on the body mass of Taiwan s bird species is scanty (Lee et al., 1998). Current body mass data are scattered in various forms, i.e., local reports and articles (e.g., Hachisuka and Udagawa, 1950, 1951), banding research data possessed by private organizations (e.g., Shiu, 1996), and museum collections. It is difficult to compile and pool the data with various qualities from different sources. In addition to local documents and reports, useful information is also available from review studies on birds. Dunning (1993) compiled body mass data of 6283 bird species primarily by reviewing published literature. The body mass data for over 60 resident species in Taiwan were presented in his work. However, the records were not obtained from the Taiwan specimens. Since body mass may vary with geographic regions, it is a question whether Dunning's data is suitable to be used for Taiwanese ecological studies on birds. The purposes of this paper are to compile the body size measurements from different local sources, and to evaluate the adequacy of applying Dunning's data in representing the bird species in Taiwan. The results generated by this study regarding body sizes will contribute to community, ecosystem and landscape studies on birds in Taiwan in the future. MATERIALS AND METHODS Data sources We employed body measurements as well as species lists from banding studies conducted between 1987 and 1995 by Wild Bird Society Taiwan. A total of 223 bird species with measurements of body mass, lengths of body, head, tail, bill and tarsus, and maximum wing span were collected. We excluded exotic birds and species captured in remote islands. All measurements were taken from living birds. Data analyses We performed an initial descriptive statistics, including mean, standard deviation, minimum, and maximum values. Outliers that showed significant deviations from the means were ignored. Among the 223 species, 138 species has sample size less than 20, and were excluded. Therefore, body mass, and lengths of bill and tarsus were analyzed for the remaining 85 species. We included bill and tarsus lengths because bill length is an important parameter for fruit-feeding species, and tarsus length is important for migratory shorebirds which use shallow water and mudflat habitats. All calculations for 223 species are available on the internet ( In addition, we evaluated the body mass data of Taiwanese birds recorded in Dunning (1993). A t-test was performed to estimate the differences of the body mass for each species in our database and Dunning s (1993). Of the 64 Taiwanese species with body mass information from Dunning (1993), ten has enough sample sizes for the analyses. Statistical analyses were performed by using SYSTAT for Windows version 9 (SPSS, 1999). RESULTS Body mass and lengths of bill and tarsus were documented for a total of 85 Taiwanese species (Tables 1 and 2), including 42 resident species, 36 winter migratory species (transient and visitor), two summer migratory species, and five species with combined status.

3 82 TAIWANIA Vol. 50, No. 2 Among the ten species of the 64 species from Dunning (1993) (Table 1), eight showed significant differences (seven cases at p < and one case at p < 0.05). Only two species showed no significant difference. A qualitative comparison between these two datasets indicated that most of these differences were rather small for smaller-sized species. As the body mass increased, there tended to be a wider difference. Table 1. Body mass measurements (g) for 85 species captured in banding research in Taiwan from 1987 to Chinese name Scientific name Sex 1) Measurement in Taiwan Data from CRC handbook 2) t-test 3) Status in Taiwan 4) N Mean SD N Mean SD 雁鴨科 Anatidae 小水鴨 Anas crecca B/U W F M 三趾鶉科 Turnicidae 棕三趾鶉 Turnix suscitator B/U R F M 鬚鴷科 Megalaimidae 五色鳥 Megalaima oorti U R 翠鳥科 Alcedinidae 翠鳥 Alcedo atthis B/U * R F M 鴟鴞科 Strigidae 黃嘴角鴞 Otus spilocephalus U R 鳩鴿科 Columbidae 斑頸鳩 Streptopelia chinensis U *** R 紅鳩 Streptopelia tranquebarica B/U R F M 鷸科 Scolopacidae 針尾鷸 Gallinago stenura B/U W 中地鷸 Gallinago megala U W 田鷸 Gallinago gallinago U W F M 中杓鷸 Numenius phaeopus U T/W F M 赤足鷸 Tringa totanus B/U T/W 小青足鷸 Tringa stagnatilis B/U T/W 青足鷸 Tringa nebularia B/U W 鷹斑鷸 Tringa glareola U T/W F M 反嘴鷸 Tringa cinerea B/U T 磯鷸 Tringa hypoleucos B/U W 黃足鷸 Tringa brevipes B/U T

4 June, 2005 Shiu et al.: Avian database in Taiwan 83 翻石鷸 Arenaria interpres B/U T/W F M 姥鷸 Calidris tenuirostris B/U T/W 漂鷸 Calidris canutus U T F M 穉鷸 Calidris ruficollis U T/W F M 丹氏穉鷸 Calidris temminckii B/U *** T 雲雀鷸 Calidris subminuta B/U T 尖尾鷸 Calidris acuminata U T F M 濱鷸 Calidris alpina U W F M 滸鷸 Calidris ferruginea B/U *** T/W F M 寬嘴鷸 Limicola falcinellus U T 紅領瓣足鷸 Phalaropus lobatus B/U T 彩鷸科 Rostratulidae F M 彩鷸 Rostratula benghalensis B/U R/W 鴴科 Charadriidae F M 金斑鴴 Pluvialis fulva B/U W 灰斑鴴 Pluvialis squatarola U NS W 小環頸鴴 Charadrius dubius B/U R/W 東方環頸鴴 Charadrius alexandrinus B/U *** R/T/W F M 蒙古鴴 Charadrius mongolus B/U T/W F M 鐵嘴鴴 Charadrius leschenaultii B/U T/W 鷗科 Laridae F M 小燕鷗 Sterna albifrons B/U R/W 鷺科 Ardeidae 小白鷺 Egretta garzetta U R 黃頭鷺 Bubulcus ibis U S/R 夜鷺 Nycticorax nycticorax B/U R 伯勞科 Laniidae

5 84 TAIWANIA Vol. 50, No. 2 紅尾伯勞 Lanius cristatus B/U T/W 鴉科 Corvidae 大卷尾 Dicrurus macrocercus U R 黑枕藍鶲 Hypothymis azurea B/U R 鶲科 Muscicapidae F M 白腹鶇 Turdus pallidus B/U W F M 赤腹鶇 Turdus chrysolaus B/U W F M 黃胸青鶲 Ficedula hyperythra B/U R F M 野鴝 Luscinia calliope B/U W F M 白眉林鴝 Tarsiger indicus B/U R F M 栗背林鴝 Tarsiger johnstoniae B/U R F M 白尾鴝 Cinclidium leucurum B/U R 山雀科 Paridae F M 青背山雀 Parus monticolus U R 燕科 Hirundinidae F M 棕沙燕 Riparia paludicola U *** R 家燕 Hirundo rustica U S/T F M 洋燕 Hirundo tahitica U R 赤腰燕 Hirundo striolata U R 鵯科 Pycnonotidae F 白頭翁 Pycnonotus sinensis U R 紅嘴黑鵯 Hypsipetes leucocephalus U R 鷦鶯科 Cisticolidae 錦鴝 Cisticola juncidis U R F M 灰頭鷦鶯 Prinia flaviventris U R M 褐頭鷦鶯 Prinia inornata U R

6 June, 2005 Shiu et al.: Avian database in Taiwan 85 繡眼科 Zosteropidae 綠繡眼 Zosterops japonicus U R 鶯科 Sylviidae 短翅樹鶯 Cettia canturians U T/W 北蝗鶯 Locustella ochotensis U T 大葦鶯 Acrocephalus arundinaceus U W 棕面鶯 Abroscopus albogularis U R 金翼白眉 Garrulax morrisonianus U R 藪鳥 Liocichla steerii U R 小彎嘴畫眉 Pomatorhinus ruficollis U R F M 山紅頭 Stachyris ruficeps U R 紋翼畫眉 Actinodura morrisoniana U R 灰頭花翼畫眉 Alcippe cinereiceps U R M 頭烏線 Alcippe brunnea U R M 繡眼畫眉 Alcippe morrisonia B/U NS R 白耳畫眉 Heterophasia auricularis U R 冠羽畫眉 Yuhina brunneiceps U R 粉紅鸚嘴 Paradoxornis webbianus U *** R 黃羽鸚嘴 Paradoxornis verreauxi U R 文鳥科 Passeridae 麻雀 Passer montanus B/U *** R 白鶺鴒 Motacilla alba U R/W M. alba alba U M. alba personata B 黃鶺鴒 Motacilla flava U W M. flava beema B M. flava flava B M. flava tschutschensis F M 白腰文鳥 Lonchura striata U R 斑文鳥 Lonchura punctulata B/U R 雀科 Fringillidae 酒紅朱雀 Carpodacus vinaceus B/U R F M 灰鷽 Pyrrhula erythaca B/U R F M 黑臉鵐 Emberiza spodocephala B/U W F M ) B = data for both sexes grouped into a single mean, F = female, M = male, U = unknown. 2) Dunning, Jr., J. B. (ed.) CRC handbook of avian body masses. CRC Press, Boca Raton, USA. 3) *p < 0.05, ***p < 0.001, NS = non-significant. 4) R = resident, S = summer visitor, T = transient, W = winter visitor.

7 86 TAIWANIA Vol. 50, No. 2 Table 2. Length of bills and tarsus (in mm) for 85 species collected in the banding research in Taiwan from 1987 to Chinese name Scientific name Sex 1) Bill length (mm) Tarsus length (mm) N Mean SD N Mean SD 雁鴨科 Anatidae 小水鴨 Anas crecca B/U F M 三趾鶉科 Turnicidae 棕三趾鶉 Turnix suscitator B/U F M 鬚鴷科 Megalaimidae 五色鳥 Megalaima oorti U 翠鳥科 Alcedinidae 翠鳥 Alcedo atthis B/U F M 鴟鴞科 Strigidae 黃嘴角鴞 Otus spilocephalus U 鳩鴿科 Columbidae 斑頸鳩 Streptopelia chinensis U 紅鳩 Streptopelia tranquebarica B/U F M 鷸科 Scolopacidae 針尾鷸 Gallinago stenura U 中地鷸 Gallinago megala U 田鷸 Gallinago gallinago U 中杓鷸 Numenius phaeopus U 赤足鷸 Tringa tetanus U 小青足鷸 Tringa stagnatilis U 青足鷸 Tringa nebularia U 鷹斑鷸 Tringa glareola U 反嘴鷸 Tringa cinerea U 磯鷸 Tringa hypoleucos U 黃足鷸 Tringa brevipes U 翻石鷸 Arenaria interpres B/U F M 姥鷸 Calidris tenuirostris U 漂鷸 Calidris canutus U 穉鷸 Calidris ruficollis U 丹氏穉鷸 Calidris temminckii U 雲雀鷸 Calidris subminuta U 尖尾鷸 Calidris acuminate U 濱鷸 Calidris alpine U 滸鷸 Calidris ferruginea B/U

8 June, 2005 Shiu et al.: Avian database in Taiwan 87 F M 寬嘴鷸 Limicola falcinellus U 紅領瓣足鷸 Phalaropus lobatus B/U F M 彩鷸科 Rostratulidae 彩鷸 Rostratula benghalensis B/U F M 鴴科 Charadriidae 金斑鴴 Pluvialis fulva U 灰斑鴴 Pluvialis squatarola U 小環頸鴴 Charadrius dubius U 東方環頸鴴 Charadrius alexandrinus B/U F M 蒙古鴴 Charadrius mongolus B/U F M 鐵嘴鴴 Charadrius leschenaultia B/U F M 鷗科 Laridae 小燕鷗 Sterna albifrons U 鷺科 Ardeidae 小白鷺 Egretta garzetta U 黃頭鷺 Bubulcus ibis U 夜鷺 Nycticorax nycticorax U 伯勞科 Laniidae 紅尾伯勞 Lanius cristatus U 鴉科 Corvidae 大卷尾 Dicrurus macrocercus U 黑枕藍鶲 Hypothymis azurea B/U F M 鶲科 Muscicapidae 白腹鶇 Turdus pallidus B/U F M 赤腹鶇 Turdus chrysolaus B/U F M 黃胸青鶲 Ficedula hyperythra B/U F M 野鴝 Luscinia calliope B/U F M

9 88 TAIWANIA Vol. 50, No. 2 白眉林鴝 Tarsiger indicus B/U F M 栗背林鴝 Tarsiger johnstoniae B/U F M 白尾鴝 Cinclidium leucurum B/U F M 山雀科 Paridae 青背山雀 Parus monticolus U 燕科 Hirundinidae 棕沙燕 Riparia paludicola U 家燕 Hirundo rustica U 洋燕 Hirundo tahitica U 赤腰燕 Hirundo striolata U 鵯科 Pycnonotidae 白頭翁 Pycnonotus sinensis U 紅嘴黑鵯 Hypsipetes leucocephalus U 鷦鶯科 Cisticolidae 錦鴝 Cisticola juncidis U 灰頭鷦鶯 Prinia flaviventris U 褐頭鷦鶯 Prinia inornata U 繡眼科 Zosteropidae 綠繡眼 Zosterops japonicus U 鶯科 Sylviidae 短翅樹鶯 Cettia canturians U 北蝗鶯 Locustella ochotensis U 大葦鶯 Acrocephalus arundinaceus U 棕面鶯 Abroscopus albogularis U 金翼白眉 Garrulax morrisonianus U 藪鳥 Liocichla steerii U 小彎嘴畫眉 Pomatorhinus ruficollis U 山紅頭 Stachyris ruficeps U 紋翼畫眉 Actinodura morrisoniana U 灰頭花翼畫眉 Alcippe cinereiceps U 頭烏線 Alcippe brunnea U 繡眼畫眉 Alcippe morrisonia U 白耳畫眉 Heterophasia auricularis U 冠羽畫眉 Yuhina brunneiceps U 粉紅鸚嘴 Paradoxornis webbianus U 黃羽鸚嘴 Paradoxornis verreauxi U 文鳥科 Passeridae 麻雀 Passer montanus U 白鶺鴒 Motacilla alba U 黃鶺鴒 Motacilla flava U 白腰文鳥 Lonchura striata U 斑文鳥 Lonchura punctulata U 雀科 Fringillidae

10 June, 2005 Shiu et al.: Avian database in Taiwan 89 酒紅朱雀 Carpodacus vinaceus B/U F M 灰鷽 Pyrrhula erythaca B/U F M 黑臉鵐 Emberiza spodocephala B/U F M ) B = data for both sexes lumped into a single mean, F = female, M = male, U = unknown. DISCUSSION The significant differences between our data and Dunning's data for two of the ten species suggests that combining body mass across geographic areas may result in inaccurate interpretations. It has been shown that avian body mass varies according to the time of a day, season, and sex (Clark, 1979), and many bird species show a variation of body sizes across geographic ranges (Dunning, 1993). These variations may reduce the adequacy of using data sets from different geographical origins to generate ecological models, such as species richness patterns. In addition, it may result in problems regarding accuracy of using few measurements from Dunning (1993), since three-fourths of the Taiwan resident bird data in Dunning (1993) has sample size less than 30 (Lee et al., 1998). Apparently more qualified data are in urgent demands in the future to generate better understanding and hypotheses. In addition to bill length, it may be worthwhile to incorporate the height and width of bird bodies into the ecological studies on birds. In this study, we reported bill length data that could be used to investigate ecological relationships of food and bill; however, the height and width of the body may affect movement and behaviors for species inhabit forests (Hedenström and Rosén, 2003; Fisher et al., 2004). Furthermore, we suggest that repeated measurements may provide more accurate assessments of body sizes (Rising and Somers, 1989; Piersma and Davidson, 1991) to reduce measurement errors in living birds. Compiling all the available body measurements from various local sources is a worthwhile endeavor that might contribute to ecological studies on birds. In this study, scattered information were collected and pooled (Hachisuka and Udagawa 1950, 1951; Lee et al. 1998), and these data could be and have been further employed to explore interesting ecological questions, such as the relationship between species richness and energy flux along an elevational gradient in Taiwan (Lee et al., 2004; Ding et al., 2005). ACKNOWLEDGEMENTS We are grateful for Dr. S. H. Wu for valuable suggestions on this manuscript. We also thank two anonymous reviewers for helpful comments. This banding research was funded by the Council of Agriculture to Wild Bird Society Taiwan. Our work was partially supported by National Science Council, Taiwan. We thank the members of Wild Bird Society Taiwan for taking the measurements.

11 90 TAIWANIA Vol. 50, No. 2 LITERATURE CITED Blackburn, T. M. and K. J. Gaston A critical assessment of the form of the interspecific relationship between abundance and body size in animals. J. Anim. Ecol. 66: Cassey, P. and T. M. Blackburn Body size trends in a Holocene island bird assemblage. Ecography 27: Clark, G. A, Jr Body weights of bird: a review. Condor 81: Cumming, G. S. and T. D. Havlicek Evolution, ecology, and multimodal distributions of body size. Ecosystems 5: Ding, T.-Z., H.-W. Yuan, S. Geng, Y.-S. Lin and P.-F. Lee Energy, body size, and diversity in relation to bird species richness along an elevational in Taiwan. Global Ecol. Biogeog. 14: in press. Dunning, J. B., Jr Body Masses of Birds of the World. In: Dunning, J. B. Jr. (ed.). CRC Handbook of Avian Body Masses. CRC Press, Boca Raton, USA. pp Fa, J. E. and A. Purvis Body size, diet and population density in Afrotropical forest mammals: a comparison with neotropical species. J. Anim. Ecol. 66: Finlay, B. L., G. F. Esteban and T. Fenchel Global diversity and body size. Nature 383: Fisher, R. J., Q. E. Q. E. Fletcher, C. K. R. Willis and R. M. Brigham Roost selection and roosting behavior of mail Common Nighthawks. Am. Midl. Nat. 151: Gaston, K. J., S. L. Chown and R. D. Mercer The animal species-body size distribution of Marion Island. PNAS 98: Gaston, K. J. and T. M. Blackburn Pattern and Process in Macroecology. Blackwell Publishing, London, UK. 392 pp. Gregory, R. D Biodiversity and body size pattern among British birds. Ecography 21: Griffiths, D Size, abundance, and energy use in communities. J. Appl. Ecol. 61: Hachisuka, M. and T. Udagawa Contributions to the ornithology of Taiwan, Part I. Quart. J. Taiwan Mus. 3: Hachisuka, M. and T. Udagawa Contributions to the ornithology of Taiwan, Part II. Quart. J. Taiwan Mus. 4: Hedenström, A. and M. Rosén Body frontal area in passerine birds. J. Avian Biol. 34: Holling, C. S Cross-scale morphology, geometry, and dynamics of ecosystems. Ecol. Monogr. 62: Lee, P.-F., T.-S. Ding and H.-J. Shiu Relationship between body mass and body length of resident bird species in Taiwan. Acta Zoologica Taiwanica 9: Lee, P.-F., T.-S. Ding, S. Geng and F.-H. Hsu Breeding bird species richness on gradients of elevation, primary productivity and human disturbance in Taiwan. J. Biogeog. 31: 1-8. Peters, R. H The Ecological Implications of Body Size. Cambridge University Press, Cambridge, USA. 329 pp. Piersma, T. and N. C. Davidson Confusions of mass and size. Auk 108: Rising, J. D. and K. M. Somers The measurement of overall body size in birds. Auk 106:

12 June, 2005 Shiu et al.: Avian database in Taiwan 91 Schluter, D. and R. Repasky Worldwide limitation of finch densities by food and other factors. Ecology 72: Shiu, H.-J Summary Report on the Morphological Measurements of Bird Banding Studies Performed by the Chinese Wild Bird Federation between 1987 and Wild Bird Society Taiwan, Taipei, Taiwan. 40 pp. (In Chinese) SPSS SYSTAT for Windows 9. SPSS Inc., Chicago, USA pp.

13 92 TAIWANIA Vol. 50, No. 2 台灣鳥類之形態特徵彙整與初探許皓捷 (1) 丁宗蘇 (2) 許嘉恩 (1) 林瑞興 (1,3) 葛兆年 (1,4) (1,5) 李培芬 ( 收稿日期 :2005 年 3 月 18 日 ; 接受日期 :2005 年 4 月 12 日 ) 摘要本文整理年間由中華鳥會, 執行鳥類繫放計畫, 所得之鳥類形態測量值, 建立其形態測量資料庫, 以做為台灣鳥類基本資料庫之內容共得 223 種鳥類之形態測值, 包括體重體長尾長喙長跗骨長等, 由於樣本數量之限制, 本研究僅報導樣本數量大於 20 之 85 種鳥類之體重喙長跗骨長本研究並比較 10 種鳥類形態測值與 Dunning (1993) 於其他地區測量之數值, 結果發現, 進行比較的種類中, 有 8 種呈現區域性變異綜合以上, 本研究的資料將對於鳥類群聚生態生態系和景觀生態之研究範疇有相當程度的貢獻關鍵詞 : 喙長體長地理變異台灣跗骨長 1. 國立台灣大學生態學與演化生物學研究所, 台北市 106 羅斯福路 4 段 1 號, 台灣 2. 國立台灣大學森林暨自然資源學系, 台北市 106 羅斯福路 4 段 1 號, 台灣 3. 行政院農業委員會特有生物研究保育中心, 南投縣 552 集集鎮民生東路 1 號, 台灣 4. 行政院農業委員會林業試驗所, 台北市 100 南海路 53 號, 台灣 5. 通信作者 Tel: ; Fax: ; leepf@ntu.edu.tw

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投影片 1 Young Bird 金城湖離堤外的客雅溪距離甚近, 當漲潮將堤外的土地淹沒之後, 鷸鴴科的鳥類必須找地方休息, 等待退潮之後再度飛到溼地進食, 此時的金城湖成為極佳的休息場所, 成群的高蹺鴴聚集湖中吵鬧而壯觀漲潮後飛到湖中的青