Introduction
Little is known about the East Asian Flyway Tundra Swan Cygnus columbianus bewickii population compared to the North-west European wintering population (Rees Reference Rees2006). China remains the core wintering area, particularly the Yangtze River floodplain (Barter et al. Reference Barter, Chen, Cao and Lei2004, Reference Barter, Lei and Cao2006, Rees Reference Rees2006, Cao et al. Reference Cao, Zhang, Barter and Lei2010), which supported c.81,000 swans (c.75% of the flyway population) during complete surveys in 2004 and 2005, (Cao et al. Reference Cao, Barter and Lei2008a), most at Poyang Lake in Jiangxi Province, but with important numbers in Anhui Province. It was formerly numerous in Korea, where fewer than 100 birds now occur annually (Moores Reference Moores2005); c.30,000 winter in Japan (Albertsen and Kanazawa Reference Albertsen and Kanazawa2002).
Recent counts of the most important wetlands for East Asian Tundra Swans have enabled an assessment of changes in their distribution and abundance in China since 2004/2005, at a time when the North-west European population is declining (Worden et al. Reference Worden, Cranswick, Crowe, McElwaine and Rees2006, Rees and Beekman Reference Rees and Beekman2010). We report Tundra Swan numbers at key Yangtze River floodplain wintering sites during 2003/2004–2009/10 inclusive. Here they feed on highly nutritious Vallisneria tubers accumulated in the mud during the summer growing season (Barzen et al. Reference Barzen, Engels, Burnham, Harris and Wu2009, Zhang et al. Reference Zhang, Cao, Barter, Fox, Zhao, Meng, Jiang and Zhu2010). Evidence suggests Vallisneria is declining at some key Tundra Swan sites (Fox et al. Reference Fox, Cao, Zhang, Barter, Zhao, Meng and Wang2011).
Methods
Study area
The Yangtze River floodplain wetlands are subject to summer monsoonal flooding followed by autumn/winter water level recession (Shankman and Liang Reference Shankman and Liang2003), creating c.10,500 km2 of numerous shallow, ephemeral, highly productive wetlands (He and Zhang Reference He and Zhang2001) that support abundant wintering waterbirds (Barter et al. Reference Barter, Chen, Cao and Lei2004, Reference Barter, Lei and Cao2006).
Poyang Lake and the Anhui Lakes (Figure S1 in the online Supplementary Material), are internationally important for tuber-feeding birds such as Tundra Swan, Swan Goose Anser cygnoides, Hooded Crane Grus monacha, Siberian Crane G. leucogeranus and White-naped Crane G. vipio (Cao Reference Cao2010).
Poyang Lake (3,300 km2), the largest freshwater lake in China (Wang et al. Reference Wang, Wang, Song, Melville and Mackinnon1993), includes Poyang Lake National Nature Reserve (NNR; 224 km2) and Nanjishan NNR (330 km2). The Anhui Lakes include Shengjin Lake NNR (76 km2) and the Anhui Anqing Yangtze Riverine Wetland Provincial Nature Reserve (ANR), the latter including seven important water bodies: Fengsha (22 km2), Baidang (57 km2), Caizi (167 km2), Wuchang (87 km2), Bo (175 km2), Huang (118 km2) and Daguan Lakes (148 km2).
Data sources
The Yangtze River floodplain was surveyed for waterbirds in February 2004 and February 2005 (Barter et al. Reference Barter, Chen, Cao and Lei2004, Reference Barter, Lei and Cao2006), counting all waterbirds present at each site (Cao et al. Reference Cao, Barter and Lei2008a). Shengjin Lake was also counted approximately fortnightly from October 2009 to April 2010 and all the main Anhui Lakes counted twice in winter 2009/2010 (13–20 December 2009; 5–10 February 2010), using the same techniques.
More or less simultaneous counts were available for Dongting Lake, Poyang Lake, Shengjin Lake and Shanghai in winters 2005/06–2008/09 (unpublished nature reserve monitoring data). ANR counts were made opportunistically at different periods within and between winters, complicating interpretation of the data but contributing to our knowledge of which sites were important for Tundra Swans.
Results and Discussion
Tundra Swan abundance and distribution in the Yangtze River floodplain
Tundra Swans were abundant in the Yangtze River estuary in the late 19th Century (Styan Reference Styan1891), and there and at Poyang Lake in the early 20th Century (La Touche Reference La Touche1934). Pre-1996, the Tundra Swan wintered from the Yangtze River floodplain north to Henan and east to inland Jiangsu, and coastal Shanghai and Zhejiang (Cao et al. Reference Cao, Wang, Wang and Barter2008b). In February 2004 and 2005, 90% of all Tundra Swans counted in the Yangtze River floodplain were in Jiangxi and Anhui (Table 1; Figure S1 in the online Supplementary Material) with fewer than 3,000 individuals in Hubei and Hunan provinces, and very few in Jiangsu province and Shanghai (Barter et al. Reference Barter, Chen, Cao and Lei2004, Reference Barter, Lei and Cao2006). Hence, since the mid-1990s, the Chinese winter range has contracted markedly to the middle Yangtze River floodplain.
Notes:
2003/2004, 2004/2005 and 2009/2010 counts conducted in early February; italics identify incomplete counts, (numbers were probably higher); - indicates no data available. 1–4: unpubl. data from the Anhui lakes (1 = mid-March; 2 = December–January; 3 = January–February; 4 = December-March); 5: February count (unpubl. data L. Cao); 6–9: data from GEF Siberian Crane Project (6-8 = late December–early January; 9 = 13 February 2009); 10: unpubl. data from “Impact of Poyang Lake Dam on wetlands and waterbirds” project (count on 27 Feb. 2010); 11–15: unpubl. data from Dongting Lake (February counts); 16–17: Hu et al. 2008; 18–20: unpubl. data from Chongming Dongtan National Bird Nature Reserve, most important site in Shanghai (February counts).
Numbers and distribution of Tundra Swans in Jiangxi province
Most swans in eastern China were counted in Jiangxi (Table 1), the majority at Poyang Lake (90% of the provincial total in 2003/2004, 99% in 2004/2005) where several discrete concentrations exceeded 5% of the Eastern Tundra Swan population (estimated at 92,000 birds; Wetlands International 2006), although distributions varied between years (Figure S2 in the online Supplementary Material). In winter 2005/06, the Jiangxi counts exceeded the current eastern Tundra Swan population estimate but more recently the province has held 43,000–82,000 birds (Table 1).
Tundra Swan abundance in the Anhui Lakes
Anhui Lakes supported 33–45% of the total floodplain Tundra Swans counted in February 2004 and 2005 (Table 1); Shengjin, Baidang, Fengsha, Wuchang and Daguan Lakes (Table 2) held internationally important numbers.
Note:
italics identify incomplete counts.
Although swan numbers at the Anhui Lakes have remained high, their distribution has changed. Shengjin Lake supported c.5% of the eastern Tundra Swan population in winters 2003/2004 and 2004/2005, but numbers there have declined since winter 2007/2008 (Table 2), although they are still of international importance (mean peak count of 1,743 from 2007/08 to 2009/10 inclusive). Numbers at Wuchang Lake have changed little since winter 2007/2008, whilst the smallest lakes (Baidang and Fengsha Lakes) have erratically supported much larger numbers of swans than in the 2003/2004 and 2004/2005 winters (Table 2). Caizi Lake, which normally holds < 1,000 swans, supported > 30,000 in 2007/2008.
Potential causes for changes in Tundra Swan numbers and distribution in the Yangtze River floodplain
Poyang and Anhui Lakes are now the major Chinese strongholds of Tundra Swans, with widely fluctuating numbers showing no trend since 2003/2004 (Cao Reference Cao2010). Swan numbers declined at Shengjin Lake, where other tuber-feeding species (e.g. Swan Goose and Hooded Crane; Cheng et al. Reference Cheng, Cao, Barter and Xu2009, Zhang et al. Reference Zhang, Cao, Barter, Fox, Zhao, Meng, Jiang and Zhu2010) have also decreased, but where numbers of ducks and other goose species have increased. These changes are thought to reflect the disappearance of Vallisneria since the early 2000s, perhaps the result of eutrophication (Fox et al. Reference Fox, Cao, Zhang, Barter, Zhao, Meng and Wang2011).
Tundra Swans recently aggregated at two small Anhui lakes: Baidang (57km2) and Fengsha Lakes (22km2). Baidang is the only lake now retaining extensive Vallisneria stands in Anhui, perhaps explaining its importance. According to local fishermen, formerly abundant Vallisneria has recently largely disappeared at Fengsha Lake, potentially explaining the collapse of swans and other tuber-feeding waterbirds in winter 2009/2010. Plans to exchange water in the next 1–2 years between Baidang or Caizi Lake and Chao Lake, a 755 km2 highly polluted wetland (Anon. 2008), to improve the water quality supplied to Hefei city and adjacent agricultural land in the vicinity, is likely to impact seriously on both lakes.
It is suggested that elevated nitrogen and phosphate levels in lake water have contributed to the collapse of Vallisneria at Shengjin Lake, affecting the tuber-feeding guild of wintering waterbirds there, including Tundra Swans (Fox et al. Reference Fox, Cao, Zhang, Barter, Zhao, Meng and Wang2011). It is therefore vital to understand the relationships between water level management, water quality and fishery management that affect the distribution and abundance of submerged plants in these lakes, in order to recommend ways to maintain the very high productivity of these ephemeral lake systems and the ecosystem services they supply to the human population, as well as ensuring they continue to support internationally significant numbers of swans in this increasingly important part of their wintering range.
Supplementary Materials
The supplementary materials for this article can be found online at journals.cambridge.org/bci
Acknowledgements
Richard Hearn and Thomas Heinicke assisted with winter fieldwork in 2007/08 and 2009/10, respectively; British Airways Communities and Conservation provided international flights for ECR and RH in 2008. Baz Hughes kindly commented on a draft of the text. The study was supported by the National Natural Science Foundation of China (Grant Nos. 30940010 & 31071941), Chinese Academy of Sciences Innovation Project (Grant No. KZCX2-YW-QN503), University of Science and Technology of China Graduate School Educational Innovation Base, plus a Chinese Academy of Sciences Visiting Professorship for Senior International Scientists (2009s1-42), State Administration of Foreign Experts Affairs, PRC and a Danske Universiteter travel grant to one of the authors (ADF). Thanks also to two anonymous referees for their constructive comments on the draft manuscript.