鸟类及其群落对崇西湿地生态恢复和生境重建的响应
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摘要
长江河口地区的潮滩芦苇沼泽湿地是水体和陆地间的重要生态通道,不仅具有减少潮滩冲刷和保护临近堤防以及陆域的重要作用,而且具有保护生物多样性、涵养淡水、向河口和近海水域补充营养物等不可替代的重要生态服务功能。一方面,区域经济发展日益增加土地资源的各种利用,使得潮滩湿地面积日益缩小,生态系统服务功能日趋退化丧失,另一方面区域社会、经济的可持续发展又依赖于潮滩湿地的生态服务功能,其中包括潮滩湿地的鸟类栖息地功能,妥善处理这个矛盾关系到长江口区域发展的可持续性。
本研究通过芦苇生境特化鸟类震旦鸦雀分布、食性、巢材、觅食生境选择等方面揭示震旦鸦雀对芦苇植被的依赖、探讨生境特化鸟类对芦苇湿地生态恢复的响应,进而通过在湿地林引入和生境要素配置情况下震旦鸦雀的觅食、分布、巢址选择、大杜鹃对栖木的利用、白鹭对微生境的利用等探讨芦苇潮滩鸟类对生境重建的响应,在此基础上,通过生态恢复、生境重建区域及其周边鸟类群落比较,探讨鸟类群落对生态恢复和生境重建的响应。
野外工作在2005年到2008年进行,研究概况如下：
.1.生态恢复和生境重建过程中芦苇潮滩湿地生态系统生命组分的改变
湿地林引入会降低林内光照强度,对芦苇产生荫蔽作用。高密度湿地林引入芦苇植被一年的时间内,会降低芦苇密度、增加芦苇枝高度、减少芦苇节数、增加节间长；随着荫蔽强度的增加,芦苇枝的死亡率显著增加,两年左右,高密度湿地林引入区域大部分芦苇消失。斑块状湿地林引入,荫蔽作用较弱,引入的湿地林可以和芦苇形成比较稳定的混合群落(见第三章)。湿地林引入也会影响芦苇昆虫,高密度湿地林引入一年之后,芦苇昆虫的发生率和丰度呈现下降趋势,两年以后,高密度湿地林引入将芦苇完全排除,相应的芦苇昆虫也消失(第五章)。可见较低密度或者斑块化引入湿地林到芦苇潮滩沼泽湿地可以避免或者降低对芦苇植被及其昆虫的负面作用。
生境要素配置实验实施一年以后,区域植物种类明显增加,由原来的一种(芦苇)增加到7种,群落植株高度显著降低,植株密度显著增加,生物量显著降低(见第四章)。表明生境要素配置实验实施以后,芦苇潮滩沼泽生态系统植物群落迅速发生了变化。
2.震旦鸦雀对生态恢复的响应(震旦鸦雀是芦苇生境特化鸟类)
从较大的地理尺度、中小地理尺度和局域尺度上,通过搜集文献资料和野外观测,论证震旦鸦雀对芦苇生境的依赖及其是否芦苇生境特化鸟类(第六章)。对震旦鸦雀在中国分布区生境利用情况的文献回顾表明83.3%的分布地点或者区域明确提及震旦鸦雀分布的生境类型,其中72.2%的分布地点和区域,震旦鸦雀分布在芦苇植被或者芦苇与其他植物混合植被中。在崇明岛,震旦鸦雀只出现在有芦苇分布的三种生境类型中(养殖鱼塘、林地生境和潮滩生境),这些生境或者以芦苇为主要植被(潮滩生境),或者包含有较大面积芦苇且和潮滩生境毗邻(养殖鱼塘和林地生境)。对崇西湿地三年的连续监测,在芦苇不分布的生境中没有记录到震旦鸦雀,大面积芦苇植被中震旦鸦雀密度较高,而稀疏乔木-芦苇生境和斑块芦苇生境中密度较低,密植乔木-芦苇混合生境形成早期,有震旦鸦雀分布,随着芦苇的消失,震旦鸦雀也不再出现在其中。这些结果表明,震旦鸦雀对芦苇植被具有非常强的依赖性；芦苇植被的存在是震旦鸦雀出现的必须条件；从分布上来说,震旦鸦雀是芦苇生境特化鸟类。
对比分析了30个东方大苇莺巢和36个震旦鸦雀巢的巢形态特征、巢材组成以及两种鸟的飞行和觅食相关的形态特征(第七章)。结果发现,东方大苇莺巢材多样性高于震旦鸦雀(2006年,P<0.05,2007年,P=0.07),表明震旦鸦雀具有较狭窄的巢材资源利用幅度；88%的震旦鸦雀巢材来自芦苇,而东方大苇莺只有29%的巢材来自芦苇,可见震旦鸦雀严重依赖芦苇作为巢材；震旦鸦雀主要使用新鲜芦苇组织,较难获取但是可以就地取材,而东方大苇莺主要用一些枯萎的植物组织,较易获取但需要飞行较远(芦苇植被中筑巢,茭白植被中搬运巢材)；而震旦鸦雀的喙强于东方大苇莺,东方大苇莺飞行能力强于震旦鸦雀,正是对各自巢材利用策略的适应。
通过束颈法收集震旦鸦雀和东方大苇莺育雏期间的食物,以分析两种鸟类育雏期间食性组成以及食物来源(第八章)。结果发现,震旦鸦雀食物中以同翅目种类最多,而东方大苇莺以双翅目最多；两种鸟类食物类群组成差异显著；东方大苇莺育雏食物的多样性和均匀度都高于震旦鸦雀,表明震旦鸦雀育雏食物的资源幅度比东方大苇莺狭窄；食物的飞行能力(有无翅膀)来看,99%的震旦鸦雀雏鸟食物为无翅的食物种类,而东方大苇莺84%的雏鸟食物为有翅的种类；无翅的食物种类主要躲藏在芦苇叶鞘或者茎秆内,对鸟类喙的力量要求比较高,而有翅的食物种类往往对飞行能力要求比较高；震旦鸦雀和东方大苇莺雏鸟食物的组成正反映了震旦鸦雀较强的喙和较弱的飞行能力、东方大苇莺较弱的喙和较强的飞行能力。
样方法沿随机样线收集芦苇枝,检测和分析震旦鸦雀的取食痕迹、潜在食物丰度、不同类型食物的取食频次等,研究震旦鸦雀觅食生境的选择(第九章)。结果发现,芦苇中茎表虫数量最多,分布最广,茎壁虫其次,茎内虫最少；震旦鸦雀主要取食茎表虫和茎内虫,并且更偏向于取食茎表虫。取食茎表虫主要在比较低矮的芦苇中,而取食茎内虫主要在相对较密集的芦苇中；取食茎表虫主要在茎表虫丰富的生境,而取食茎内虫主要是在茎内虫丰富的生境中。较高和较粗的芦苇枝生境中具有较多的茎表虫和茎内虫。茎表虫取食的量和芦苇枝的高度呈显著负相关,茎内虫的取食与新芦苇枝的密度显著正相关,茎壁虫的取食程度和死亡芦苇枝比例呈显著负相关,不同类型的昆虫被取食的程度和各类昆虫的分布和数量呈显著正相关。这些结果表明,昆虫的可提供性是震旦鸦雀觅食生境选择的主要影响因素,而芦苇(高度、密度、基径等)通过影响昆虫或者被昆虫影响而影响震旦鸦雀觅食生境的选择。
3.鸟类群落关键种对生境重建的响应
通过连续跟踪震旦鸦雀和棕头鸦雀在芦苇收割进程中以及收割结束后的分布格局的变化,探讨鸟类对芦苇植被斑块化的即时响应(第十章)。震旦鸦雀对芦苇的移除和斑块化反应很快,随着芦苇收割的进行,震旦鸦雀减少了对相应区域的利用,而对棕头鸦雀受影响不明显；收割结束1周以后,震旦鸦雀开始逐步恢复对收割区域剩余斑块芦苇的利用,四周左右震旦鸦雀出现在所有收割剩余斑块芦苇区域,但是种群数量、集群规模均偏小,而棕头鸦雀受芦苇收割影响较小,收割过程以及收割后均出现在整个区域。这些结果暗示,生境泛化的棕头鸦雀受生境重建过程中生境斑块化的影响要小于震旦鸦雀；生境特化在一定程度上限制了震旦鸦雀生境利用的可塑性,然则经过一段时间后,震旦鸦雀对生境斑块化有适应的趋势。
在芦苇收割前后通过样方法研究了芦苇收割对震旦鸦雀觅食分布的影响(第十一章)。研究表明,芦苇收割显著减少了震旦鸦雀潜在食物资源,改变了震旦鸦雀觅食分布并使震旦鸦雀提前转移到新生芦苇枝上取食,芦苇收割后残留老芦苇枝对震旦鸦雀取食具有重要影响。芦苇收割时保留一定的斑块状和条带状老芦苇有利于震旦鸦雀对食物资源的利用。
湿地林引入芦苇沼泽以后,由于荫蔽作用,芦苇密度下降、昆虫丰度也降低(第三和第五章),比较了湿地林引入区域和没有湿地林引入区域芦苇枝上震旦鸦雀食物和取食痕迹的情况(第十二章)。发现,湿地林引入短时间内,昆虫被取食的数量显著下降,远低于未引入湿地林的区域；一年的时间内,昆虫取食比例在引入湿地林和未引入湿地林的芦苇沼泽中均下降,但是引入湿地林的芦苇沼泽中下降更大,表明湿地林引入芦苇沼泽以后影响了鸟类对芦苇昆虫的利用。这些结果表明,作为对湿地林引入的响应,震旦鸦雀的觅食分布发生了相应的变化,震旦鸦雀较少出现在湿地林引入区域觅食。
系统搜寻纯芦苇生境、密植乔木-芦苇混合生境、斑块芦苇生境以及防护林带中震旦鸦雀的巢,研究震旦鸦雀巢址选择对生境重建的响应(第十三章)。2006年到2008年共找到117个震旦鸦雀的巢,99%的巢分布在芦苇中,其中86%分布在纯芦苇生境中；密植乔木-芦苇混合生境中,只找到一个震旦鸦雀巢；纯芦苇生境中震旦鸦雀巢密度显著高于斑块芦苇植被；有巢和无巢芦苇样方间,绝大部分芦苇和昆虫相关的参数差异不显著。这些结果表明,湿地林引入和芦苇植被的斑块化都会降低区域作为震旦鸦雀繁殖生境的适宜性,而在纯芦苇生境中震旦鸦雀对巢址的选择性不明显,也即只要是较大面积的芦苇植被即可以成为其巢址。
比较了2006年到2008年震旦鸦雀在崇西湿地纯芦苇生境、斑块芦苇生境、密植乔木-芦苇混合植被以及稀疏乔木-芦苇混合植被中的分布情况以及集群规模的差异(第十四章)。发现,纯芦苇生境中震旦鸦雀密度显著高于斑块芦苇生境、密植乔木-芦苇生境和稀疏乔木-芦苇生境。密植乔木-芦苇生境中,2007年芦苇植被消失,此后再也没有记录到震旦鸦雀；而稀疏乔木-芦苇混合生境中,震旦鸦雀一直有记录,而且密度与斑块芦苇生境差异不显著；集群规模较大的非繁殖季节,各生境类型(密植乔木-芦苇混合生境、纯芦苇生境和斑块芦苇生境)中各集群规模(1-2只／集群,3-7只／集群和7只以上集群)出现频次相近,而繁殖季节,1-2只／集群出现频次最高,7只以上集群很少见,且在各生境类型间出现频次相似。这些结果表明,震旦鸦雀对湿地林引入和生境要素配置产生的芦苇斑块化在分布上作出了响应,这种分布格局的形成与各生境食物资源变化以及震旦鸦雀巢址分布在各生境间的差异有一定关系。
通过增加设置人工栖木,比较大杜鹃在增加栖木的芦苇沼泽和未增加栖木的芦苇沼泽中的出现频次、行为组成、巢寄生的差异(第十五章)。发现栖木的增加(湿地林引入),可以增加区域大杜鹃的丰度和出现频次,相应的也改变了大杜鹃的行为组成,在增加栖木的区域,大杜鹃有较多的栖停行为,而未增加栖木的区域,大杜鹃的飞行行为较多；增加栖木的区域,大杜鹃飞入芦苇中的次数要显著的高于未增加栖木的区域,栖木的增加便于大杜鹃观察和搜寻区域寄主,因而也增加了区域大杜鹃主要寄主-东方大苇莺被巢寄生的风险。这些结果表明,湿地林引入会增加区域的栖木,而大杜鹃通过分布和行为对栖木的增加做出响应,进而可能影响其繁殖成功率。
野外观测白鹭对生境要素配置区微生境(水线区域、裸地、水面和芦苇捆)的利用及其行为组成(第十六章)。白鹭在生境要素配置区的休息行为频次最高,而所有行为出现在水线附近的频次最高；表明生境要素配置区作为白鹭休息场所似乎要甚于作为觅食场所；而如果考虑各微生境的相对面积,则白鹭对芦苇捆的偏好更加明显,表明白鹭对生境要素配置区栖停条件有要求。45%的觅食行为发生在水线附近；40%的休息行为发生在芦苇捆上。生境要素配置区毗邻生境的可利用性对白鹭是否利用生境要素配置区有决定性的影响,周边没有被潮水淹没时(周边生境可提供性高),生境要素配置区白鹭数量也较高,相应的周边生境被潮水淹没时,利用生境要素配置区的白鹭数量较低。可见,生境要素配置区所处的位置、周边区域鹭类的数量是影响生境要素配置区鹭类实际生境利用的重要因素；生境要素配置区鹭类的数量随着周边鹭类的数量变动而变动,潮汐涨落及其对生境要素配置区域的淹没状态对鹭类利用生境要素配置区有重要影响。
4.鸟类群落对生态恢复和生境重建的响应
采用BACI (Before-After-Control-Impact)方法,比较湿地林引入区域和芦苇沼泽区域冬季鸟类群落,以及1年后冬季鸟类群落的变化(第十七章)。湿地林引入芦苇植被短时间内,整个区域冬季鸟类密度和物种数量都下降,而多样性指数有所上升；对于芦苇沼泽鸟类有负面的影响,密度降低、物种丰富度和多样性指数降低。而湿地林内部,鸟类多样性上升,但是密度下降。表明湿地林引入对鸟类群落的影响是双刃剑,对原有鸟类群落产生负面作用,也为新的鸟类群落形成提供了机遇。
比较了湿地林引入区域和生境要素配置实施区域与原有的纯芦苇植被、光滩生境中鸟类群落的差异,同时也比较了生态恢复和生境重建区域与临近周边地区鸟类群落的差异,以探讨鸟类群落对湿地生态恢复和生境重建的响应(第十八章)。湿地林生境中鸟类群落的物种丰富度最高；生境要素配置区总的鸟类丰富度和湿地鸟类丰富度与芦苇、光滩生境相近。生态恢复和生境重建区域,总的鸟类丰富度和湿地鸟类丰富度都要高于周边邻近区域,但是丰度接近或者低于周边邻近区域,湿地鸟类中,迁徙鸟、留鸟和冬候鸟丰度要高于周边区域。
对崇西湿地不同生境类型中鸟类群落同资源种团进行了分析(第十九章)。林泽区域鸟类群落物种丰富度、同资源种团数量均是最高的；林泽区域集合了其他区域生境特征的同时也成为较多鸟类适宜的栖息地；生境要素配置实验区鸟类群落物种丰富度和同资源种团数量均不高,但是物种多样性和同资源种团多样性均是最高的。从湿地依赖性同资源种团的分析来看,湿地专性物种同资源种团无论种类还是数量在芦苇沼泽、光滩和生境要素配置实验区鸟类群落中所占的比例都要远高于林泽；而生境要素配置区鸟类群落中依赖湿地和不依赖湿地的鸟类种类和数量比例组成差不多,表明该区域栖息条件对湿地和非湿地鸟类都比较适宜(第十八和十九章)。
The reed(Phragmites australis)-dominated tidal marshes at the Yangtze River Estuary are important buffer zone between the water and the land, as they can not only reduce erosion and protect the nearby dike, but also provide other ecosystem services, such as protecting biodiversity, water conservation, nutrition sources to estuarine and coastal waters. On the one hand, the tidal marsh wetland has suffered decrease in area and degradation in ecosystem functions because of reclamations and disturbance from the near society, and on the other hand, the nearby society needs the tidal marsh to provide critical ecosystem service, such as serving as habitat of wildlife (given that the wetlands of Yangtze River estuary are important staging habitat of shorebird following the Australian-Asia flyway). A solution to this conflict is important for sustainable development of the Yangtze River delta.
In this thesis, I have performed field-experiments and observations to reveal the responses of birds and avian community to ecological restoration and bird habitat restoration in Phragmites-dominated tidal marshes of the Chongxi Wetland Research Center. We have performed ecological restoration project first and then have selected and performed two treatments as habitat restoration:construction of forested wetland in reed-dominated tidal marsh by inducing trees that may adapt wetland area (CFW) and reallocation of habitat elements in reed-dominated tidal marsh by eradicating reeds, changing topography, and constructed water area (RHE). I studied the distribution, diets, nest materials and feeding habitat selection of reed habitat specialist-reed parrotbill, Paradoxornis heudei. Through changes of feeding behavior, distribution, and nest site selection of reed parrotbill, use of perch site by the common cuckoo, and micro-habitat use of egret, I studied the responses of birds inhabitating in tidal marsh to the bird habitat restoration. Through comparing the structure, diversity and guilds of bird communities between restored tidal marshes and control ones, I studied the responses of bird community to the ecological restoration and bird habitat restoration conducted at the Phragmites-dominated tidal marsh. Field works have been performed in 2005 to 2007 and main results are below.
1. changes of living component to the ecological restoration and bird habitat restoration
Construction of forested wetland in reedbed reduced the illumination in the area, thus casued shading to the reed shoots. In the first year, in area with high density of forest, the density of reed shoots decreased, the height of reed shoots increased, and the number of internodes decreased. The mortality of reed shoots inseased with the density of forest. In the second year after forested, the reed shoots dispeared in area with high density. In patched forested area, the trees and the reed shoots coexisted (Chapter 3). In forested reedbed, the abundance of the insects on reed shoots decreased and dispeared when the reed shoots dispeared (Chapter 5). It seems the introduction of low density forest or patches of trees can reduce the potential negative effects to the reedbed and insects on them.
Through reallocation of habitat elements in reed-dominated tidal marsh, we conducted bird habitat restoration. One year after reallocation of habitat elements, the diversity of plant increased. The height of plant community decreased. The plant density and biomass decreased (chapter 4).
2. Response of Reed Parrotbill to ecological restoration
Through reviewing papers and field observations, I tried to demonstrate whether the Reed Parrotbill is a reed habitat specialist (Chapter 6). The results showed that in its distribution range in China,83.3% of sites where Reed Parrotbills had been witnessed were reed vegetation or habitat with Common Reeds. In Chongming Island, Reed Parrotbills were observed only in habitats with reeds, such as fish ponds, woodland and tidal marshes. This habitat was either reed-dominated habitat or a habitat with reed patches and near those dominated by reeds. In the local scale, during three-year continuous monitoring, Reed Parrotbills has not been observed in habitats without reeds. The density of Reed Parrotbill was significant higher in reedbeds than in reeds with dense trees and patches of reeds. In the habitat of reeds with dense trees, Reed Parrotbills have not been seen in this habitat since the reeds disappeard in the second year. These results indicate that Reed Parrotbill strongly depend on reed vegetation as its habitat and the Reed Parrotbill is a reed-habitat specialist.
I compared the characteristics, material composition and morphology of nests of Oriental Great Reed Warblers (OGW) and Reed Parrotbills (RP), and related these parameters with their flight- and foraging-related morphologies (Chapter 7). The diversity of nest material was higher for OGW than for RP (2006, P<0.05,2007, p=0.07), indicating a narrower resource rage used by RP than OGW.88% of RP's nest material was tissues of the Common Reed and 81% of OGW's nest material was tissues of the Wild Rice. RPs used mainly tissues from living reeds, which need a strong beak to get, while OGWs used mainly dead plant tissues, which are easy to get, but need flying a distance to transport. The RP had stronger beak than OGW, while OGW had strong flight ability than RP, which is adapted well to each other's nest material usage.
Through neck ligatures, we collected and compared food composition delivered to nestling by OGWs and RPs (Chapter 8). RPs used mainly insect from the order Hemiptera, while OGWs used mainly insects from the order Diptera. There was a significant difference in food composition between the OGW and the RP. The diversity of food was higher for OGWs than RPs, indicating a relative narrow food resource range of RP.99% of insects delivered to nestling by RP were wingless, while 84% of those by OGWs were winged. The wingless insects mainly hid within reed shoots, which was easier to target but hard to retrieve. A strong beak is needed. The winged insects often flied in the air, which need relative strong flight ability to pursue and get them. The food composition of RPs reflected a strong beak but weak flight ability, while that of OGWs reflected a weak beak but strong flight ability.
I studied the foraging habitat selection of RPs by collected reed samples to determine foraging marks, food resource abundance, and characteristics of reed shoots (Chapter 9). The scale insects were the most abundant food resource and the stem-boring caterpillars were the least abundant one. The RPs foraged mainly on scale insects and stem-boring caterpillars and preferred to scale insects more. The RPs foraged on scale insects in areas with short reed shoots, while they foraged on stem-boring caterpillars in areas with dense reed shoots. The amount of insects attacked by RPs was depend on the density of insects, with higher density higher attacked probability. There was a significantly negative relationship between the amount of attacked scale insects and the height of reed shoots. There was a significant positive relationship between the attacked stem-boring caterpillars and the density of newly emerged reed shoots. The predation intensity of each insect significantly positively related to the abundance of each insect. These results indicated that the food resource availability was important factors influenced the foraging habitat selection of RPs.
3. Response of bird species to habitat restoration
Through continuous monitoring the distribution of Reed Parrotbills (RP) and the Vinous-throated Parrotbills (VP) during reed-harvest, I studied the instant responses of RP and VP to reed-harvest (Chapter 10). RPs responded to the reed-harvest fast. RPs reduced their activities in areas where reed-harvest was going on. One week after reed-harvest, RPs came back to the harvested area close to bigger area of reed shoots and used the reed patches as foraging habitat. Four weeks after reed-harvest, RPs redistributed in all reed-harvest area with reed patches. But the group size and density were smaller than those before reed-harvest. The VP did not show obviously signs affected by the reed-harvest during and after harvesting. These results indicated that habitat fragmentation influenced less on a habitat generalist species (VP) than on a habitat specialist (RP), while the RP also had a relative plasticity to adapt to habitat fragmentation, though slower than a habitat generalist.
I studied the impacts of reed cutting on foraging of Reed Parrotbill from January to April 2007 (Chapter 11). Reed cutting decreased potential food resource of Reed Parrotbill significantly. Reed cutting changed the foraging habitat of Reed Parrotbill, and shifted their foraging place from dead reed shoots to growing reed shoots before they suited Reed Parrotbill to feed on them. Short time after reed cutting, the residual dead reed shoots have an important role for foraging of Reed Parrotbill. And these indicated that keeping some reed patches uncut could facilitate Reed Parrotbill foraging on growing reed shoots and lessen food shortage impacts of reed cutting.
Construction of forested wetland reduced the density of reed shoots and also the abundance of insects on reed shoots (see Chapter 3 and 5). The foraging intensity by Reed Parrotbills of insects on reed shoots in reedbed and forested wetland was compared (Chapter 12). The amount of insects that was preyed by RPs decreased significantly after forested wetland had been constructed shortly. This indicated that construction of forested wetland affected the foraging distribution of RPs.
I systematically searched reedbed, reedbed with dense trees, reed patches and protection forest for nests of the RP to study the effects of habitat restoration on nest site selection of RPs (Chapter 13).117 nests were found from 2006 to 2008.99% of these nests were found in reed vegetation and 86% was in reedbed. Only one nest was found in reedbed with dense trees. The nest density was significant higher in reedbeds than in reed patches. There were no significant difference in reed- and food-related characteristics between nest site and site without nest. These results indicated that, first, RPs were heavily depend on reed vegetation as their nesting habitat, second, construction of forested wetland and reallocation of habitat elements in reed-dominated tidal marsh could reduce the quality of these areas as RPs'nesting habitat.
The distribution and group size of RPs in 2006 to 2008 among reedbed, reed patches, reedbed with dense trees and reedbed with sparse trees was compared (Chapter 14). The density of RPs was significant higher in reedbed than in all other habitats. Since the reed shoots in reedbed with dense trees disappeared in 2007, RPs has not been seen in this habitat. There was RPs in reedbed with sparse trees in all three years, and the density in reedbed with sparse trees was not significantly different with that in reed patches. The frequencies of different group size (1-2 inds./group, 3-7inds./group and>7inds./group) in reedbed, reedbed with dense trees and reedbed with sparse trees were similar. These results indicated that RPs responded to the reed vegetation fragmentation induced by construction of forested wetland and reallocation of habitat elements in their distribution and this response was consistent with distribution in nest site and foraging intensity.
Construction of forested wetland can increase perch site for some birds. I studied the response of Common Cuckoo to a simulation of perch site increase by erecting bamboo pole in reedbeds (Chapter 15). The abundance and emerging frequency of Common Cuckoo increased with increased perch sites. Common Cuckoo showed more frequency in perching behavior in area with increased perch sites than in area without and more flying frequency in areas without perch sites. The nest parasitism rates of OGW by Common Cuckoos increased in area with perch sites. These results demonstrated that construction of forested wetland could increase perch sites, thus increase abundance of the Common Cuckoo and also change the behavior of the Common Cuckoo.
Observations were conducted to determine the use of microhabitat (waterline, waters, naked area and perch site simulated by bunch of reed shoots) in area with reallocation of habitat elements (RHE) by Litter Egret (Chapter 16). Little Egrets showed more frequency of rest behavior in RHE area and waterline was more frequently used than other microhabitats. 45% of foraging happened near waterline and 40% of resting in bunch of reed shoots. The abundance of Litter Egret in RHE areas was higher when there was more in tidal flat nearby. While the tide covered the tidal flat nearby, there was also no egret in RHE area. This indicated that usage of RHE area as habitats by egrets depended on the abundance of egrets in a larger scale.
4. Responses of avian community to ecological restoration and habitat restoration
With BACI (before-after-control-impact), the avian community in winter was compared between reedbed and forested wetland (Chapter 17). After construction of forested wetland (with dense trees), during one year, the density and species richness of the whole area decreased and the diversity increased. The density, species richness and diversity index of wintering birds depending on reedbed decreased. Within the forested wetland, the density decreased and the diversity increased. These results indicated that construction of forested wetland could show negative effects on avian community depending on habitat before habitat restoration; also provide opportunity for new avian community.
In CFW area, the avian community had the highest species richness while in RHE area, the species richness of whole community and wetland birds were similar with those in reedbeds and tidal flat. The area with CFW and RHE had higher species richness in whole community and wetland bird community than vicinal area, while the abundance was similar to the vicinal area. The abundances of migrants, residents and wintering birds belonging to wetland birds were higher than vicinal area (Chapter 18).
There was the highest species richness and number of response guilds in CFW area. In RHE area, there was the highest diversity index in species and guilds. The species and abundance of obligate wetland species were higher in RHE, reedbed and tidal flat than in CFW, While there almost the same proportion of obligate wetland species and upland species, indicating the RHE area was both good habitat for wetland birds and land birds (chapter 19).
本研究通过芦苇生境特化鸟类震旦鸦雀分布、食性、巢材、觅食生境选择等方面揭示震旦鸦雀对芦苇植被的依赖、探讨生境特化鸟类对芦苇湿地生态恢复的响应,进而通过在湿地林引入和生境要素配置情况下震旦鸦雀的觅食、分布、巢址选择、大杜鹃对栖木的利用、白鹭对微生境的利用等探讨芦苇潮滩鸟类对生境重建的响应,在此基础上,通过生态恢复、生境重建区域及其周边鸟类群落比较,探讨鸟类群落对生态恢复和生境重建的响应。
野外工作在2005年到2008年进行,研究概况如下：
.1.生态恢复和生境重建过程中芦苇潮滩湿地生态系统生命组分的改变
湿地林引入会降低林内光照强度,对芦苇产生荫蔽作用。高密度湿地林引入芦苇植被一年的时间内,会降低芦苇密度、增加芦苇枝高度、减少芦苇节数、增加节间长；随着荫蔽强度的增加,芦苇枝的死亡率显著增加,两年左右,高密度湿地林引入区域大部分芦苇消失。斑块状湿地林引入,荫蔽作用较弱,引入的湿地林可以和芦苇形成比较稳定的混合群落(见第三章)。湿地林引入也会影响芦苇昆虫,高密度湿地林引入一年之后,芦苇昆虫的发生率和丰度呈现下降趋势,两年以后,高密度湿地林引入将芦苇完全排除,相应的芦苇昆虫也消失(第五章)。可见较低密度或者斑块化引入湿地林到芦苇潮滩沼泽湿地可以避免或者降低对芦苇植被及其昆虫的负面作用。
生境要素配置实验实施一年以后,区域植物种类明显增加,由原来的一种(芦苇)增加到7种,群落植株高度显著降低,植株密度显著增加,生物量显著降低(见第四章)。表明生境要素配置实验实施以后,芦苇潮滩沼泽生态系统植物群落迅速发生了变化。
2.震旦鸦雀对生态恢复的响应(震旦鸦雀是芦苇生境特化鸟类)
从较大的地理尺度、中小地理尺度和局域尺度上,通过搜集文献资料和野外观测,论证震旦鸦雀对芦苇生境的依赖及其是否芦苇生境特化鸟类(第六章)。对震旦鸦雀在中国分布区生境利用情况的文献回顾表明83.3%的分布地点或者区域明确提及震旦鸦雀分布的生境类型,其中72.2%的分布地点和区域,震旦鸦雀分布在芦苇植被或者芦苇与其他植物混合植被中。在崇明岛,震旦鸦雀只出现在有芦苇分布的三种生境类型中(养殖鱼塘、林地生境和潮滩生境),这些生境或者以芦苇为主要植被(潮滩生境),或者包含有较大面积芦苇且和潮滩生境毗邻(养殖鱼塘和林地生境)。对崇西湿地三年的连续监测,在芦苇不分布的生境中没有记录到震旦鸦雀,大面积芦苇植被中震旦鸦雀密度较高,而稀疏乔木-芦苇生境和斑块芦苇生境中密度较低,密植乔木-芦苇混合生境形成早期,有震旦鸦雀分布,随着芦苇的消失,震旦鸦雀也不再出现在其中。这些结果表明,震旦鸦雀对芦苇植被具有非常强的依赖性；芦苇植被的存在是震旦鸦雀出现的必须条件；从分布上来说,震旦鸦雀是芦苇生境特化鸟类。
对比分析了30个东方大苇莺巢和36个震旦鸦雀巢的巢形态特征、巢材组成以及两种鸟的飞行和觅食相关的形态特征(第七章)。结果发现,东方大苇莺巢材多样性高于震旦鸦雀(2006年,P<0.05,2007年,P=0.07),表明震旦鸦雀具有较狭窄的巢材资源利用幅度；88%的震旦鸦雀巢材来自芦苇,而东方大苇莺只有29%的巢材来自芦苇,可见震旦鸦雀严重依赖芦苇作为巢材；震旦鸦雀主要使用新鲜芦苇组织,较难获取但是可以就地取材,而东方大苇莺主要用一些枯萎的植物组织,较易获取但需要飞行较远(芦苇植被中筑巢,茭白植被中搬运巢材)；而震旦鸦雀的喙强于东方大苇莺,东方大苇莺飞行能力强于震旦鸦雀,正是对各自巢材利用策略的适应。
通过束颈法收集震旦鸦雀和东方大苇莺育雏期间的食物,以分析两种鸟类育雏期间食性组成以及食物来源(第八章)。结果发现,震旦鸦雀食物中以同翅目种类最多,而东方大苇莺以双翅目最多；两种鸟类食物类群组成差异显著；东方大苇莺育雏食物的多样性和均匀度都高于震旦鸦雀,表明震旦鸦雀育雏食物的资源幅度比东方大苇莺狭窄；食物的飞行能力(有无翅膀)来看,99%的震旦鸦雀雏鸟食物为无翅的食物种类,而东方大苇莺84%的雏鸟食物为有翅的种类；无翅的食物种类主要躲藏在芦苇叶鞘或者茎秆内,对鸟类喙的力量要求比较高,而有翅的食物种类往往对飞行能力要求比较高；震旦鸦雀和东方大苇莺雏鸟食物的组成正反映了震旦鸦雀较强的喙和较弱的飞行能力、东方大苇莺较弱的喙和较强的飞行能力。
样方法沿随机样线收集芦苇枝,检测和分析震旦鸦雀的取食痕迹、潜在食物丰度、不同类型食物的取食频次等,研究震旦鸦雀觅食生境的选择(第九章)。结果发现,芦苇中茎表虫数量最多,分布最广,茎壁虫其次,茎内虫最少；震旦鸦雀主要取食茎表虫和茎内虫,并且更偏向于取食茎表虫。取食茎表虫主要在比较低矮的芦苇中,而取食茎内虫主要在相对较密集的芦苇中；取食茎表虫主要在茎表虫丰富的生境,而取食茎内虫主要是在茎内虫丰富的生境中。较高和较粗的芦苇枝生境中具有较多的茎表虫和茎内虫。茎表虫取食的量和芦苇枝的高度呈显著负相关,茎内虫的取食与新芦苇枝的密度显著正相关,茎壁虫的取食程度和死亡芦苇枝比例呈显著负相关,不同类型的昆虫被取食的程度和各类昆虫的分布和数量呈显著正相关。这些结果表明,昆虫的可提供性是震旦鸦雀觅食生境选择的主要影响因素,而芦苇(高度、密度、基径等)通过影响昆虫或者被昆虫影响而影响震旦鸦雀觅食生境的选择。
3.鸟类群落关键种对生境重建的响应
通过连续跟踪震旦鸦雀和棕头鸦雀在芦苇收割进程中以及收割结束后的分布格局的变化,探讨鸟类对芦苇植被斑块化的即时响应(第十章)。震旦鸦雀对芦苇的移除和斑块化反应很快,随着芦苇收割的进行,震旦鸦雀减少了对相应区域的利用,而对棕头鸦雀受影响不明显；收割结束1周以后,震旦鸦雀开始逐步恢复对收割区域剩余斑块芦苇的利用,四周左右震旦鸦雀出现在所有收割剩余斑块芦苇区域,但是种群数量、集群规模均偏小,而棕头鸦雀受芦苇收割影响较小,收割过程以及收割后均出现在整个区域。这些结果暗示,生境泛化的棕头鸦雀受生境重建过程中生境斑块化的影响要小于震旦鸦雀；生境特化在一定程度上限制了震旦鸦雀生境利用的可塑性,然则经过一段时间后,震旦鸦雀对生境斑块化有适应的趋势。
在芦苇收割前后通过样方法研究了芦苇收割对震旦鸦雀觅食分布的影响(第十一章)。研究表明,芦苇收割显著减少了震旦鸦雀潜在食物资源,改变了震旦鸦雀觅食分布并使震旦鸦雀提前转移到新生芦苇枝上取食,芦苇收割后残留老芦苇枝对震旦鸦雀取食具有重要影响。芦苇收割时保留一定的斑块状和条带状老芦苇有利于震旦鸦雀对食物资源的利用。
湿地林引入芦苇沼泽以后,由于荫蔽作用,芦苇密度下降、昆虫丰度也降低(第三和第五章),比较了湿地林引入区域和没有湿地林引入区域芦苇枝上震旦鸦雀食物和取食痕迹的情况(第十二章)。发现,湿地林引入短时间内,昆虫被取食的数量显著下降,远低于未引入湿地林的区域；一年的时间内,昆虫取食比例在引入湿地林和未引入湿地林的芦苇沼泽中均下降,但是引入湿地林的芦苇沼泽中下降更大,表明湿地林引入芦苇沼泽以后影响了鸟类对芦苇昆虫的利用。这些结果表明,作为对湿地林引入的响应,震旦鸦雀的觅食分布发生了相应的变化,震旦鸦雀较少出现在湿地林引入区域觅食。
系统搜寻纯芦苇生境、密植乔木-芦苇混合生境、斑块芦苇生境以及防护林带中震旦鸦雀的巢,研究震旦鸦雀巢址选择对生境重建的响应(第十三章)。2006年到2008年共找到117个震旦鸦雀的巢,99%的巢分布在芦苇中,其中86%分布在纯芦苇生境中；密植乔木-芦苇混合生境中,只找到一个震旦鸦雀巢；纯芦苇生境中震旦鸦雀巢密度显著高于斑块芦苇植被；有巢和无巢芦苇样方间,绝大部分芦苇和昆虫相关的参数差异不显著。这些结果表明,湿地林引入和芦苇植被的斑块化都会降低区域作为震旦鸦雀繁殖生境的适宜性,而在纯芦苇生境中震旦鸦雀对巢址的选择性不明显,也即只要是较大面积的芦苇植被即可以成为其巢址。
比较了2006年到2008年震旦鸦雀在崇西湿地纯芦苇生境、斑块芦苇生境、密植乔木-芦苇混合植被以及稀疏乔木-芦苇混合植被中的分布情况以及集群规模的差异(第十四章)。发现,纯芦苇生境中震旦鸦雀密度显著高于斑块芦苇生境、密植乔木-芦苇生境和稀疏乔木-芦苇生境。密植乔木-芦苇生境中,2007年芦苇植被消失,此后再也没有记录到震旦鸦雀；而稀疏乔木-芦苇混合生境中,震旦鸦雀一直有记录,而且密度与斑块芦苇生境差异不显著；集群规模较大的非繁殖季节,各生境类型(密植乔木-芦苇混合生境、纯芦苇生境和斑块芦苇生境)中各集群规模(1-2只／集群,3-7只／集群和7只以上集群)出现频次相近,而繁殖季节,1-2只／集群出现频次最高,7只以上集群很少见,且在各生境类型间出现频次相似。这些结果表明,震旦鸦雀对湿地林引入和生境要素配置产生的芦苇斑块化在分布上作出了响应,这种分布格局的形成与各生境食物资源变化以及震旦鸦雀巢址分布在各生境间的差异有一定关系。
通过增加设置人工栖木,比较大杜鹃在增加栖木的芦苇沼泽和未增加栖木的芦苇沼泽中的出现频次、行为组成、巢寄生的差异(第十五章)。发现栖木的增加(湿地林引入),可以增加区域大杜鹃的丰度和出现频次,相应的也改变了大杜鹃的行为组成,在增加栖木的区域,大杜鹃有较多的栖停行为,而未增加栖木的区域,大杜鹃的飞行行为较多；增加栖木的区域,大杜鹃飞入芦苇中的次数要显著的高于未增加栖木的区域,栖木的增加便于大杜鹃观察和搜寻区域寄主,因而也增加了区域大杜鹃主要寄主-东方大苇莺被巢寄生的风险。这些结果表明,湿地林引入会增加区域的栖木,而大杜鹃通过分布和行为对栖木的增加做出响应,进而可能影响其繁殖成功率。
野外观测白鹭对生境要素配置区微生境(水线区域、裸地、水面和芦苇捆)的利用及其行为组成(第十六章)。白鹭在生境要素配置区的休息行为频次最高,而所有行为出现在水线附近的频次最高；表明生境要素配置区作为白鹭休息场所似乎要甚于作为觅食场所；而如果考虑各微生境的相对面积,则白鹭对芦苇捆的偏好更加明显,表明白鹭对生境要素配置区栖停条件有要求。45%的觅食行为发生在水线附近；40%的休息行为发生在芦苇捆上。生境要素配置区毗邻生境的可利用性对白鹭是否利用生境要素配置区有决定性的影响,周边没有被潮水淹没时(周边生境可提供性高),生境要素配置区白鹭数量也较高,相应的周边生境被潮水淹没时,利用生境要素配置区的白鹭数量较低。可见,生境要素配置区所处的位置、周边区域鹭类的数量是影响生境要素配置区鹭类实际生境利用的重要因素；生境要素配置区鹭类的数量随着周边鹭类的数量变动而变动,潮汐涨落及其对生境要素配置区域的淹没状态对鹭类利用生境要素配置区有重要影响。
4.鸟类群落对生态恢复和生境重建的响应
采用BACI (Before-After-Control-Impact)方法,比较湿地林引入区域和芦苇沼泽区域冬季鸟类群落,以及1年后冬季鸟类群落的变化(第十七章)。湿地林引入芦苇植被短时间内,整个区域冬季鸟类密度和物种数量都下降,而多样性指数有所上升；对于芦苇沼泽鸟类有负面的影响,密度降低、物种丰富度和多样性指数降低。而湿地林内部,鸟类多样性上升,但是密度下降。表明湿地林引入对鸟类群落的影响是双刃剑,对原有鸟类群落产生负面作用,也为新的鸟类群落形成提供了机遇。
比较了湿地林引入区域和生境要素配置实施区域与原有的纯芦苇植被、光滩生境中鸟类群落的差异,同时也比较了生态恢复和生境重建区域与临近周边地区鸟类群落的差异,以探讨鸟类群落对湿地生态恢复和生境重建的响应(第十八章)。湿地林生境中鸟类群落的物种丰富度最高；生境要素配置区总的鸟类丰富度和湿地鸟类丰富度与芦苇、光滩生境相近。生态恢复和生境重建区域,总的鸟类丰富度和湿地鸟类丰富度都要高于周边邻近区域,但是丰度接近或者低于周边邻近区域,湿地鸟类中,迁徙鸟、留鸟和冬候鸟丰度要高于周边区域。
对崇西湿地不同生境类型中鸟类群落同资源种团进行了分析(第十九章)。林泽区域鸟类群落物种丰富度、同资源种团数量均是最高的；林泽区域集合了其他区域生境特征的同时也成为较多鸟类适宜的栖息地；生境要素配置实验区鸟类群落物种丰富度和同资源种团数量均不高,但是物种多样性和同资源种团多样性均是最高的。从湿地依赖性同资源种团的分析来看,湿地专性物种同资源种团无论种类还是数量在芦苇沼泽、光滩和生境要素配置实验区鸟类群落中所占的比例都要远高于林泽；而生境要素配置区鸟类群落中依赖湿地和不依赖湿地的鸟类种类和数量比例组成差不多,表明该区域栖息条件对湿地和非湿地鸟类都比较适宜(第十八和十九章)。
The reed(Phragmites australis)-dominated tidal marshes at the Yangtze River Estuary are important buffer zone between the water and the land, as they can not only reduce erosion and protect the nearby dike, but also provide other ecosystem services, such as protecting biodiversity, water conservation, nutrition sources to estuarine and coastal waters. On the one hand, the tidal marsh wetland has suffered decrease in area and degradation in ecosystem functions because of reclamations and disturbance from the near society, and on the other hand, the nearby society needs the tidal marsh to provide critical ecosystem service, such as serving as habitat of wildlife (given that the wetlands of Yangtze River estuary are important staging habitat of shorebird following the Australian-Asia flyway). A solution to this conflict is important for sustainable development of the Yangtze River delta.
In this thesis, I have performed field-experiments and observations to reveal the responses of birds and avian community to ecological restoration and bird habitat restoration in Phragmites-dominated tidal marshes of the Chongxi Wetland Research Center. We have performed ecological restoration project first and then have selected and performed two treatments as habitat restoration:construction of forested wetland in reed-dominated tidal marsh by inducing trees that may adapt wetland area (CFW) and reallocation of habitat elements in reed-dominated tidal marsh by eradicating reeds, changing topography, and constructed water area (RHE). I studied the distribution, diets, nest materials and feeding habitat selection of reed habitat specialist-reed parrotbill, Paradoxornis heudei. Through changes of feeding behavior, distribution, and nest site selection of reed parrotbill, use of perch site by the common cuckoo, and micro-habitat use of egret, I studied the responses of birds inhabitating in tidal marsh to the bird habitat restoration. Through comparing the structure, diversity and guilds of bird communities between restored tidal marshes and control ones, I studied the responses of bird community to the ecological restoration and bird habitat restoration conducted at the Phragmites-dominated tidal marsh. Field works have been performed in 2005 to 2007 and main results are below.
1. changes of living component to the ecological restoration and bird habitat restoration
Construction of forested wetland in reedbed reduced the illumination in the area, thus casued shading to the reed shoots. In the first year, in area with high density of forest, the density of reed shoots decreased, the height of reed shoots increased, and the number of internodes decreased. The mortality of reed shoots inseased with the density of forest. In the second year after forested, the reed shoots dispeared in area with high density. In patched forested area, the trees and the reed shoots coexisted (Chapter 3). In forested reedbed, the abundance of the insects on reed shoots decreased and dispeared when the reed shoots dispeared (Chapter 5). It seems the introduction of low density forest or patches of trees can reduce the potential negative effects to the reedbed and insects on them.
Through reallocation of habitat elements in reed-dominated tidal marsh, we conducted bird habitat restoration. One year after reallocation of habitat elements, the diversity of plant increased. The height of plant community decreased. The plant density and biomass decreased (chapter 4).
2. Response of Reed Parrotbill to ecological restoration
Through reviewing papers and field observations, I tried to demonstrate whether the Reed Parrotbill is a reed habitat specialist (Chapter 6). The results showed that in its distribution range in China,83.3% of sites where Reed Parrotbills had been witnessed were reed vegetation or habitat with Common Reeds. In Chongming Island, Reed Parrotbills were observed only in habitats with reeds, such as fish ponds, woodland and tidal marshes. This habitat was either reed-dominated habitat or a habitat with reed patches and near those dominated by reeds. In the local scale, during three-year continuous monitoring, Reed Parrotbills has not been observed in habitats without reeds. The density of Reed Parrotbill was significant higher in reedbeds than in reeds with dense trees and patches of reeds. In the habitat of reeds with dense trees, Reed Parrotbills have not been seen in this habitat since the reeds disappeard in the second year. These results indicate that Reed Parrotbill strongly depend on reed vegetation as its habitat and the Reed Parrotbill is a reed-habitat specialist.
I compared the characteristics, material composition and morphology of nests of Oriental Great Reed Warblers (OGW) and Reed Parrotbills (RP), and related these parameters with their flight- and foraging-related morphologies (Chapter 7). The diversity of nest material was higher for OGW than for RP (2006, P<0.05,2007, p=0.07), indicating a narrower resource rage used by RP than OGW.88% of RP's nest material was tissues of the Common Reed and 81% of OGW's nest material was tissues of the Wild Rice. RPs used mainly tissues from living reeds, which need a strong beak to get, while OGWs used mainly dead plant tissues, which are easy to get, but need flying a distance to transport. The RP had stronger beak than OGW, while OGW had strong flight ability than RP, which is adapted well to each other's nest material usage.
Through neck ligatures, we collected and compared food composition delivered to nestling by OGWs and RPs (Chapter 8). RPs used mainly insect from the order Hemiptera, while OGWs used mainly insects from the order Diptera. There was a significant difference in food composition between the OGW and the RP. The diversity of food was higher for OGWs than RPs, indicating a relative narrow food resource range of RP.99% of insects delivered to nestling by RP were wingless, while 84% of those by OGWs were winged. The wingless insects mainly hid within reed shoots, which was easier to target but hard to retrieve. A strong beak is needed. The winged insects often flied in the air, which need relative strong flight ability to pursue and get them. The food composition of RPs reflected a strong beak but weak flight ability, while that of OGWs reflected a weak beak but strong flight ability.
I studied the foraging habitat selection of RPs by collected reed samples to determine foraging marks, food resource abundance, and characteristics of reed shoots (Chapter 9). The scale insects were the most abundant food resource and the stem-boring caterpillars were the least abundant one. The RPs foraged mainly on scale insects and stem-boring caterpillars and preferred to scale insects more. The RPs foraged on scale insects in areas with short reed shoots, while they foraged on stem-boring caterpillars in areas with dense reed shoots. The amount of insects attacked by RPs was depend on the density of insects, with higher density higher attacked probability. There was a significantly negative relationship between the amount of attacked scale insects and the height of reed shoots. There was a significant positive relationship between the attacked stem-boring caterpillars and the density of newly emerged reed shoots. The predation intensity of each insect significantly positively related to the abundance of each insect. These results indicated that the food resource availability was important factors influenced the foraging habitat selection of RPs.
3. Response of bird species to habitat restoration
Through continuous monitoring the distribution of Reed Parrotbills (RP) and the Vinous-throated Parrotbills (VP) during reed-harvest, I studied the instant responses of RP and VP to reed-harvest (Chapter 10). RPs responded to the reed-harvest fast. RPs reduced their activities in areas where reed-harvest was going on. One week after reed-harvest, RPs came back to the harvested area close to bigger area of reed shoots and used the reed patches as foraging habitat. Four weeks after reed-harvest, RPs redistributed in all reed-harvest area with reed patches. But the group size and density were smaller than those before reed-harvest. The VP did not show obviously signs affected by the reed-harvest during and after harvesting. These results indicated that habitat fragmentation influenced less on a habitat generalist species (VP) than on a habitat specialist (RP), while the RP also had a relative plasticity to adapt to habitat fragmentation, though slower than a habitat generalist.
I studied the impacts of reed cutting on foraging of Reed Parrotbill from January to April 2007 (Chapter 11). Reed cutting decreased potential food resource of Reed Parrotbill significantly. Reed cutting changed the foraging habitat of Reed Parrotbill, and shifted their foraging place from dead reed shoots to growing reed shoots before they suited Reed Parrotbill to feed on them. Short time after reed cutting, the residual dead reed shoots have an important role for foraging of Reed Parrotbill. And these indicated that keeping some reed patches uncut could facilitate Reed Parrotbill foraging on growing reed shoots and lessen food shortage impacts of reed cutting.
Construction of forested wetland reduced the density of reed shoots and also the abundance of insects on reed shoots (see Chapter 3 and 5). The foraging intensity by Reed Parrotbills of insects on reed shoots in reedbed and forested wetland was compared (Chapter 12). The amount of insects that was preyed by RPs decreased significantly after forested wetland had been constructed shortly. This indicated that construction of forested wetland affected the foraging distribution of RPs.
I systematically searched reedbed, reedbed with dense trees, reed patches and protection forest for nests of the RP to study the effects of habitat restoration on nest site selection of RPs (Chapter 13).117 nests were found from 2006 to 2008.99% of these nests were found in reed vegetation and 86% was in reedbed. Only one nest was found in reedbed with dense trees. The nest density was significant higher in reedbeds than in reed patches. There were no significant difference in reed- and food-related characteristics between nest site and site without nest. These results indicated that, first, RPs were heavily depend on reed vegetation as their nesting habitat, second, construction of forested wetland and reallocation of habitat elements in reed-dominated tidal marsh could reduce the quality of these areas as RPs'nesting habitat.
The distribution and group size of RPs in 2006 to 2008 among reedbed, reed patches, reedbed with dense trees and reedbed with sparse trees was compared (Chapter 14). The density of RPs was significant higher in reedbed than in all other habitats. Since the reed shoots in reedbed with dense trees disappeared in 2007, RPs has not been seen in this habitat. There was RPs in reedbed with sparse trees in all three years, and the density in reedbed with sparse trees was not significantly different with that in reed patches. The frequencies of different group size (1-2 inds./group, 3-7inds./group and>7inds./group) in reedbed, reedbed with dense trees and reedbed with sparse trees were similar. These results indicated that RPs responded to the reed vegetation fragmentation induced by construction of forested wetland and reallocation of habitat elements in their distribution and this response was consistent with distribution in nest site and foraging intensity.
Construction of forested wetland can increase perch site for some birds. I studied the response of Common Cuckoo to a simulation of perch site increase by erecting bamboo pole in reedbeds (Chapter 15). The abundance and emerging frequency of Common Cuckoo increased with increased perch sites. Common Cuckoo showed more frequency in perching behavior in area with increased perch sites than in area without and more flying frequency in areas without perch sites. The nest parasitism rates of OGW by Common Cuckoos increased in area with perch sites. These results demonstrated that construction of forested wetland could increase perch sites, thus increase abundance of the Common Cuckoo and also change the behavior of the Common Cuckoo.
Observations were conducted to determine the use of microhabitat (waterline, waters, naked area and perch site simulated by bunch of reed shoots) in area with reallocation of habitat elements (RHE) by Litter Egret (Chapter 16). Little Egrets showed more frequency of rest behavior in RHE area and waterline was more frequently used than other microhabitats. 45% of foraging happened near waterline and 40% of resting in bunch of reed shoots. The abundance of Litter Egret in RHE areas was higher when there was more in tidal flat nearby. While the tide covered the tidal flat nearby, there was also no egret in RHE area. This indicated that usage of RHE area as habitats by egrets depended on the abundance of egrets in a larger scale.
4. Responses of avian community to ecological restoration and habitat restoration
With BACI (before-after-control-impact), the avian community in winter was compared between reedbed and forested wetland (Chapter 17). After construction of forested wetland (with dense trees), during one year, the density and species richness of the whole area decreased and the diversity increased. The density, species richness and diversity index of wintering birds depending on reedbed decreased. Within the forested wetland, the density decreased and the diversity increased. These results indicated that construction of forested wetland could show negative effects on avian community depending on habitat before habitat restoration; also provide opportunity for new avian community.
In CFW area, the avian community had the highest species richness while in RHE area, the species richness of whole community and wetland birds were similar with those in reedbeds and tidal flat. The area with CFW and RHE had higher species richness in whole community and wetland bird community than vicinal area, while the abundance was similar to the vicinal area. The abundances of migrants, residents and wintering birds belonging to wetland birds were higher than vicinal area (Chapter 18).
There was the highest species richness and number of response guilds in CFW area. In RHE area, there was the highest diversity index in species and guilds. The species and abundance of obligate wetland species were higher in RHE, reedbed and tidal flat than in CFW, While there almost the same proportion of obligate wetland species and upland species, indicating the RHE area was both good habitat for wetland birds and land birds (chapter 19).
引文
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