随着天然滨海湿地正以惊人的速度消失，东亚—澳大利西亚迁徙路线上多种水鸟的数 量急剧下降。作为天然湿地被改造后出现的生境类型，人工湿地为部分水鸟的生存和繁衍 提供了替代性的栖息地。准确地评估人工生境在缓解天然湿地快速消失后对水鸟多样性维 持所发挥的作用，是本条迁徙路线保护工作的一项重要内容。海岸盐池是滨海地区常见的 一种人工湿地类型，在世界范围内被认为是一种能够支持大量水鸟的功能性湿地。位于渤 海湾北部的南堡盐池区（总面积 290 km2）是世界上最大的盐池区之一，有关该盐池在水鸟 多样性维持方面的作用尚缺乏专项研究。为此，我们于 2013 - 2016 年对该盐池区及其毗邻 的天然潮间带滩涂的水鸟开展了系统的野外调查，分析和评估了该湿地对于东亚—澳大利 西亚迁徙路线水鸟生存和繁衍的生态功能和潜在价值，并对未来该湿地的保护和管理工作 提出了建议。本论文的主要结果如下: 1. 调查期间在南堡盐池区共记录到水鸟 89 种，占我国水鸟物种总数的 31.37%，其中 27 种水鸟的种群数量超过其在东亚—澳大利西亚迁徙路线总种群数量的 1%，优势物种和 常见物种包括弯嘴滨鹬（Calidris ferruginea）（最大数量占该迁徙路线种群数量的 69%）、 红腹滨鹬（Calidris canutus）（占 40%）、黑翅长脚鹬（Himantopus himantopus）（占 61%）、 反嘴鹬（Recurvirostra avosetta）（占 57%）、鹤鹬（Tringa erythropus）（占 54%）、黑尾塍鹬 （Limosa limosa）（占 11%）和遗鸥（Larus relictu）（占 43%）等。在该盐池区中，包括取 食和休息的个体在内，高潮阶段春季和秋季记录到的最大水鸟数量分别是 96,000 只和 93,500 只；而相应地低潮阶段在滩涂上记录到的最大水鸟数量分别为 73,000 只和 20,000 只。在春季北迁时期，该地区利用滩涂的水鸟的平均数量较利用盐池的多；而秋季南迁期 间则刚好相反，盐池中的鸟类数量相对更多。冬季该地区利用盐池的鸟类非常少，但有数 千只个体在滩涂上取食。从湿地功能上看，不管是高潮阶段还是低潮阶段，春季水鸟在内 陆盐池区（中心位置距潮间带距离约 8.7 km）主要是取食，在海边盐池区（两处中心位置 距潮间带距离分别约为 1.0 km 和 2.8 km）主要是休息。从各种水鸟对盐池的利用程度看， 黑尾塍鹬、反嘴鹬、泽鹬（Tringa stagnatilis）和黑翅长脚鹬等水鸟几乎只利用盐池，而红 腹滨鹬、大滨鹬（Calidris tenuirostris）、斑尾塍鹬（Limosa lapponica）、白腰杓鹬（Numenius arquata）、遗鸥和灰斑鸻（Pluvialis squatarola）等更偏好滩涂湿地。鉴于南堡盐池区以及毗 邻的潮间带滩涂是东亚—澳大利西亚迁徙水鸟的关键中停地，该地区正面临着巨大的开发 压力，因此尽早建立涵盖滩涂和盐池的自然保护区是加强该湿地及其水鸟保护的一条最有 效途径。 2. 作为水鸟的重要栖息地，南堡盐池区的底栖动物丰富，主要以卤虫（Artemia spp.）、摇蚊幼虫（摇蚊科）以及盐水蝇的幼虫、蛹（水蝇科）构成。经鉴定，水蝇科和摇蚊科的 主要物种分别为盐生摇蚊（Chironomus salinarius）和蓝额水蝇（Ephydra glauca）。在我们 采集的所有样品中，卤虫、摇蚊幼虫以及盐水蝇的幼虫、蛹和成体所占的数量比例分别为 29%、27%、17%、26%和 0.3%。无灰干重（Ash Free Dry Weight, AFDW）生物量的比例 （不包括其他稀少的底栖动物）分别为 8.4%、8.3%、16.0%、66.0%和 0.7%。底栖动物分 布不均匀，密度在各盐池间差异悬殊。卤虫，摇蚊幼虫以及盐水蝇的幼虫、蛹和成体的平 均密度分别为 700 ± 208、619 ± 194、445 ± 140、852 ± 289 和 7 ± 3 个/m2（means ± SE）， 相应的平均生物量为 0.110 ± 0.037、0.095 ± 0.032、0.240 ± 0.004、1.098 ± 0.374、0.009 ± 0.004 g AFDW /m2。底栖动物中盐水蝇蛹的生物量最大。盐池周边的盐水蝇成体密度和生 物量分别为 11,890 ± 3,963 个/m2 和 15.219 ± 5.073g AFDW /m2。广义线性混合模型分析表 明，在春季，随着季节的推进，底栖动物的密度逐渐上升，而水深、盐度、水质情况以及 位置对底栖动物的分布与密度存在重要影响。研究发现，南堡盐池区高密度的底栖动物为 东亚—澳大利西亚迁徙路线上的候鸟提供了充足的食物资源。 3.使用广义线性混合模型分析表明，三个类别的水鸟（小型鸻鹬类，非小型鸻鹬类及 非鸻鹬类水鸟）对南堡盐池区的取食受水深、盐度、水质、人为干扰、天敌及日期等多方 面因素的影响。盐度、水质、人为干扰和天敌对水鸟取食数量的影响在三个类别间是类似 的，而水深和日期的影响则因类别而异。三类水鸟的取食数量都随着盐度的上升而减少， 但模型中盐度只显著影响了非鸻鹬类水鸟的取食数量。三类水鸟在无废水的盐池中取食数 量均显著大于有废水的盐池。天敌和人为干扰的存在会减少水鸟的取食数量，但在三个类 别中均没有达到显著的影响。非小型鸻鹬类取食数量随水深的增加而减少；小型鸻鹬类和 非鸻鹬类水鸟的取食数量则随水深增加而呈二次效应，但小型鸻鹬类的二次效应不显著。 鸻鹬类最适宜的取食水深为 0 -10 cm，非鸻鹬类水鸟的最适宜取食水深为 30 – 40 cm。日 期对小型鸻鹬类有显著的二次效应，对非小型鸻鹬类的二次效应不显著。随着迁徙季节推 进非小型鸻鹬类取食数量持续减少，小型鸻鹬类和非鸻鹬类水鸟都在迁徙季节的中期取食 数量最多。未发现其他显著影响水鸟取食的因素。除了 2014 年的小型和非小型鸻鹬类， 2016 的小型鸻鹬类，三类水鸟研究期间在各盐池间的数量都存在显著差异。 4. 在本研究区，泽鹬、反嘴鹬、黑翅长脚鹬主要利用盐池生境，黑腹滨鹬（Calidris alpina）、弯嘴滨鹬、红颈滨鹬（Calidris ruficollis）、环颈鸻（Charadrius alexandrinus）、尖 尾滨鹬（Calidris acuminata）则综合利用滩涂和盐池生境，翻石鹬（Arenaria interpres）、翘 嘴鹬（Xenus cinereus）则基本只利用滩涂生境。采用视频记录手段（或者直接观察取食行 为）为主，查看部分死亡个体的胃容物分析为辅的研究方法，我们对南堡湿地的鸻鹬类食 性进行了研究，并对泽鹬和黑尾塍鹬的取食成功速率进行了分析，对取食效率进行估算。 结果发现：鸻鹬类在南堡湿地的食性组成多样化，包括贝类、螺类、甲壳类、双翅目、鞘 翅目和蛛形目等；鸻鹬类的食性组成与其对滩涂和盐池生境的利用模式较吻合，来自盐池的主要食物为卤虫、盐水蝇幼虫、蛹和成体以及摇蚊幼虫等，不同鸟类的食性存在差异； 泽鹬和黑尾塍鹬的取食成功速率分别为 0.259 ± 0.009/s（n = 211）和 0.223 ± 0.015/s（n = 96）(means ± SE)；泽鹬和黑尾塍鹬在盐池中的取食效率（单位时间无灰干重摄入）分别为 0.167 ± 0.011 mg AFDW /s 和 0.172 ± 0.005 mg AFDW /s。对比每日摄入能量与每日能量需 求（daily energy requirement，DER），我们推测泽鹬需在夜间取食，黑尾塍鹬应另有其他更 高取食效率的取食地。主要在盐池中取食的鸻鹬类水鸟相对的肌胃干重和相对盐腺干重都 要小于滩涂上以贝类或螺类为食的物种，而两种生境混合利用的水鸟相对的肌胃干重和相 对盐腺干重的大小则介于上述两者之间。 5. 反嘴鹬是在南堡盐池区取食和繁殖的一种常见候鸟。2015 - 2016 年我们对其繁殖表 现进行了为期两年的调查与监测。结果发现，两年之中其巢表观存活率分别为 52.7%和 47.9%，每日存活率分别为 0.984 和 0.970，以孵化期 24 天的平均存活率分别为 67.4%和 48.2%。南堡盐池区反嘴鹬的巢存活率与世界其他地区同类的巢存活率相当，较高的存活率 得益于该地区盐池较低的人为干扰。造成繁殖失败的主要原因是弃巢，而捕食者的影响因 素相对较低。MARK 模型分析显示，巢存活率与卵数、巢区位置、巢距地面高度、巢龄、 繁殖时间、温度和降雨等因素相关，各因素对存活率的影响在两年的两个研究地有所不同。 此研究表明，在人为干扰较低的情况下，盐场可以成为反嘴鹬的一个理想繁殖地。

The natural coastal wetlands of East Asian-Australasian Flyway (EAAF) are disappearing at alarming rates, leading to rapid declines of many populations of waterbirds in this most speciesrich flyway in the world. Converted from natural wetlands, artificial wetlands provide alternatively habitat for the surviving and breeding of many waterbirds species. Identification and assessment of possible alternative habitats that may buffer the loss of natural wetlands and support the waterbirds diversity should be one of priority objects in this flyway. Coastal saltpans are functional wetlands that support large numbers of waterbirds worldwide. Nanpu Saltpans on the northern Bohai Bay in China, is one of the largest (290 km2 ) saltpan complexes in the world, but it’s still unclear about its value for maintaining waterbirds diversity. In this thesis, we document the value of Nanpu Saltpans for supporting waterbirds in the East Asian-Australasian Flyway. The research was carried out from 2013 to 2016, focused waterbird both in the saltpans and on the adjacent natural intertidal mudflats (57 km2 ). We analyzed and evaluated the ecological function and potential value of the Nanpu Saltpans for the survival and reproduction of waterbirds in the East Asian-Australasian Flyway, and provided suggestions about wetland protection and management in the future. The main results of the research are as follows: 1: During research period, total 89 waterbird species were recorded, 27 had maximum numbers exceeded the 1% threshold value of known flyway populations. These include a large numbers of Curlew Sandpiper Calidris ferruginea (69% of the EAAF population), Red Knot Calidris canutus (40%), Black-winged Stilt Himantopus himantopus (61%), Pied Avocet Recurvirostra avosetta (57%), Spotted Redshank Tringa erythropus (54%), Black-tailed Godwit Limosa limosa (11%), and Relict Gull Larus relictus (43%). The maximum numbers of waterbirds in spring and autumn in Nanpu Saltpans were 96,000 and 93,500, respectively. These peak numbers included both foraging and resting birds. While the peak numbers on the adjacent mudflats at low tide amounted to 73,000 and 20,000 waterbirds in spring and autumn, respectively, and almost of them were foraging birds. In winter, few birds fed in saltpans, but several thousand fed on mudflats. Waterbirds used inland ponds (8.7 km from center to the intertidal area) mainly for feeding both during low tide and high tide, and used the nearshore ponds (two sites, 1.0 km and 2.8 km from center to the intertidal area respectively) mainly for high tide roosting. Some species such as Black-tailed Godwits, Pied Avocets, Marsh Sandpipers Tringa stagnatilis and Black-winged Stilts occurred mainly in saltpans; other species preferred mudflats, such as Red Knots, Great Knots Calidris tenuirostris, Bar-tailed Godwits Limosa lapponica, Eurasian Curlews Numenius arquata, Relict Gulls, and Grey Plovers Pluvialis squatarola. The Nanpu Saltpan complex with the adjacent intertidal mudflats is a key staging area for waterbirds in the East AsianAustralasian Flyway and this area is facing enormous development pressures, so establish a nature reserve as soon as possible is the most effective way to strengthen the protection of the wetlands and waterbirds. 2. As an important habitat for waterbirdes, the benthic fauna of Nanpu Saltpans is rich, mainly composed of brine shrimp (Artemia spp.), larvae of chironomid (Chironomidae), larvae and pupa of brine fly (Ephydridae). Chironomus salinarius and Ephydra glauca were the most important species of the brine fly and the chironomid respectively. In all of the samples we collected, the number of brine shrimp, larvae of chironomid, larvae, pupa and adult of brine fly accounted for 29%, 27%, 17%, 26% and 0.3%, respectively. The proportion of biomass (ash free dry weight, AFDW) of these benthic fauna (excluding other rare benthic animals) were 8.4%, 8.3%, 16%, 66% and 0.7%, respectively. There were great differences of the benthic fauna density among ponds. The mean density of brine shrimp, chironomid larvae, brine fly larvae, pupa and adult were 700 ± 208, 619 ± 194, 445 ± 140, 852 ± 289, and 7 ± 3 /m2 (means ± SE), respectively, and the corresponding biomass were 0.110 ± 0.037, 0.095 ± 0.032, 0.240 ± 0.004, 1.098 ± 0.374, and 0.009 ± 0.004 g AFDW /m2 . The biomass of the brine fly pupa was the largest in benthic fauna. The density and biomass of brine fly adult on the ground surrounding saltpans were 11890 ± 3963 /m2 and 15.219 ± 5.073 g AFDW /m2 , respectively. The generalized linear mixed models (GLMM) showed, in the spring, as the times goes on, the density of benthic gradually increased, and the water depth, salinity, sampling location and water quality strongly affected the distribution and density of benthic fauna. This study found that high density of benthic fauna in Nanpu Saltpans can provide ample food for migratory waterbirds in East Asian-Australasian Flyway. 3. Using the generalized linear mixed models (GLMM) showed that three groups of waterbirds (small size shorebirds, non-small size shorebirds, and non-shorebirds waterbirds) used Nanpu Saltpans for feeding were affected by many factors such as water depth, salinity, water quality, human disturbance, predators and date. The effects of salinity, water quality, human disturbance and predators on the number of waterbirds feeding in saltpans are similar among three groups, while the effects of water depth and date varied among groups. The number of three groups of waterbirds feeding in saltpans decreased with the increase of salinity, but only the nonshorebirds were significantly affected by salinity in models. The feeding number of three groups of waterbirds were significant greater in ponds without pollution than those polluted. The presence of predators and human disturb would reduce the feeding number of waterbirds, but no significant effect were found among three groups. The feeding number of non-small size shorebirds decreased as the water depth increased; while the feeding number of small-size shorebirds and non shorebirds waterbirds changed with quadratic effect as water depth increased, however, the quadratic effect of small-size shorebirds was not significant. The optimum feeding depth were 0 - 10 cm and 30 - 40 cm for shorebirds and non-shorebirds waterbirds respectively. The quadratic effect of the date on the small size shorebirds was significant, but not on the non-small size shorebirds. In spring, as the time goes on, the feeding number of non-small size shorebirds decreased; while the feeding number of small size shorebirds and non-shorebirds were the highest in the middle of spring migration season. No other factors were found significantly affect the number of waterbirds feeding. The number of three groups of waterbirds feeding were all significantly different among ponds, except the small size and non-small size shorebirds in 2014, and small size shorebirds in 2016. 4. In Nanpu wetlands, Marsh Sandpiper, Pied Avocet, Black-winged Stilt mainly used saltpans; Dunlin Calidris alpina, Curlew Sandpiper, Red-necked Stint Calidris ruficollis, Kentish Plover Charadrius alexandrines and Sharp-tailed Sandpiper Calidris acuminata use both saltpans and intertidal mudflats; Ruddy Turnstone Arenaria interpres and Terek Sandpiper Xenus cinereus mainly mainly used mudflats. By analysis of video records of shorebirds feeding (or directly observer feeding behaviors) and supplemented by checking gizzard contents of some dead individuals, we studied diet of the shorebirds in Nanpu wetlands, and analyzed feeding rate and intake rate of Marsh Sandpiper and Black-tailed Godwit in saltpans. We found that the diet of shorebirds in Nanpu wetlands were diversity, including bivalve, gastropod, Diptera, Coleoptera, Araneae et al. The diet of different species of shorebirds were matched with their habitat use pattern between saltpans and mudflats in study area. Diet from saltpans were mainly brine shrimp, chironomid larvae, brine fly larvae, pupa and adult, and they were different among shorebird species. The feeding rates of Marsh Sandpiper and Black-tailed Godwit were 0.259 ± 0.009/s (n = 211) and 0.223 ± 0.015/s (n = 96) (means ± SE), respectively. The intake rate of Marsh Sandpiper and Black-tailed Godwit were 0.167 ± 0.011 mg AFDW /s and 0.172 ± 0.005 mg AFDW /s, respectively. Compared daily energy intake with daily energy requirement (DER), we predicted Marsh Sandpiper needed feed at night, and Black-tailed Godwit had other higher intake rate feeding sites. The relative dry weight of gizzard and salt gland, both lighter in the saltpan used species than intertidal mudflats used species, and those of mix habitats used species were between them. 5 Pied Avocet is a common migratory species feeding and breeding in Nanpu Saltpans. We monitored the nests of avocet in 2015 and 2016. The apparent nest survival rates were 52.7% and 47.9% respectively in the two years. The daily nest survival rates (DSR) were 0.984 and 0.97 respectively. The average nest survival rate (calculated from DSR with 24 incubation days) were 67.4% and 48.2% respectively. This nest survival rates in Nanpu Saltpans were comparable to other sites around the world, and the high survival rates were due to low human disturbance in saltpans. The main reasons of nest failure were nest abandoned, and the effect of predators were limited. Programme MARK models showed that the survival rates of nest were related with the number of eggs, nest site, nest height to ground, nest age, breeding date, temperature, and precipitation. The effects of most factors on survival rates were different between two years. This study showed that in the case of low human disturbance, the saltpans can be an ideal breeding ground for the Pied Avocet.