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Article

How Do Waterbird Communities Respond to Multi-Scale Environmental Variables in the Satellite Wetlands Surrounding a Ramsar Site, Shengjin Lake in China?

by
Chengrong Pan
1,†,
Sheng Xu
1,†,
Zhenbing Qian
1,
Qichen Liao
1,
Tongxinyu Wu
2 and
Guangyao Wang
2,*
1
Anhui Province Eco-Environmental Monitoring Center, Hefei 230071, China
2
School of Resources and Environmental Engineering, Anhui University, Hefei 230601, China
*
Author to whom correspondence should be addressed.
†
These authors contributed equally to this work.
Diversity 2025, 17(3), 176; https://doi.org/10.3390/d17030176
Submission received: 12 January 2025 / Revised: 8 February 2025 / Accepted: 25 February 2025 / Published: 28 February 2025
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Abstract

:
The global degradation and loss of natural wetlands are increasingly threatening wetland-dependent taxa, particularly waterbirds, which are highly vulnerable to environmental changes. In response to these threats, an increasing number of waterbirds are relocating to surrounding satellite wetlands in search of compensatory habitats. However, how waterbirds utilize these satellite wetlands and respond to varying environmental variables remain poorly understood. In the winter of 2022–2023 and summer of 2023, we conducted surveys on waterbird assemblages in 49 satellite wetlands of different types (reservoirs, aquaculture ponds, paddy fields and natural ponds) surrounding Shengjin Lake, a Ramsar site, and analyzed the relationship between community metrics and environmental factors. Large numbers of waterbirds were recorded during both summer and winter, including several threatened and nationally protected species. Species richness and number of individuals varied significantly across wetland types, with aquaculture ponds supporting the highest number of species and individuals. These two metrics showed positive correlations with wetland areas and landscape connectivity in both seasons. Species richness was also positively correlated with habitat diversity in summer. The number of individuals was positively correlated with habitat diversity and negatively with distance to human settlements, but the pattern was in contrast to that in winter. The Shannon–Wiener diversity index displayed a similar pattern among wetland types in winter but did not in summer. We detected no effects of environmental factors on the diversity index. Species composition differed markedly between wetland types in both seasons, especially between reservoirs and aquaculture ponds. To safeguard waterbird communities in the middle and lower reaches of the Yangtze River, we recommend integrating surrounding satellite wetlands into the regional wetland network and reducing human disturbances, particularly during the winter months.

1. Introduction

Global wetland ecosystems are experiencing unprecedented rates of degradation and loss, with an estimated 35% of wetlands having disappeared since the 1970s [1]. This widespread decline is primarily driven by human activities, including land conversion for agriculture, urbanization, industrial development and climate change [2]. The destruction of wetlands has severe consequences for biodiversity, particularly for species that rely on these habitats for critical life stages. Waterbirds, which are highly dependent on wetlands for breeding, migration and feeding, have been especially affected, with many species showing declines in population and distribution [3,4]. These trends underscore the urgent need for conservation efforts to protect and restore wetland habitats, not only for the benefit of wetland species but also for the continued provision of essential ecosystem services.
In the context of global wetland degradation and loss, many waterbird species inhabiting natural wetlands (hereafter “core wetland”) have to find compensatory habitats in surrounding small and micro-wetlands, which here are referred to as “satellite wetlands”. These satellite wetlands, which mainly encompass natural ponds and artificial wetlands such as aquaculture ponds, reservoirs and paddy fields, are often connected with the core wetland via various ecological processes, such as regional biogeochemical cycles. Studies have found that some waterbirds will move to these surrounding satellite wetlands when their natural habitats experience degradation or loss [5,6]. Among others, human disturbances are often more intense in these satellite wetlands, potentially threatening the waterbirds moving here [5]. Despite their importance, the ecological functions of these altered habitats remain poorly understood, necessitating further research to evaluate their role in maintaining biodiversity and supporting waterbird communities. A comprehensive understanding of how waterbirds interact with these satellite wetlands is crucial for developing effective conservation strategies and habitat management practices.
Environmental variables not only shape the structure and composition of waterbird communities but also play a significant role in determining the suitability of satellite wetlands for different species [7,8]. The utilization of satellite wetlands by waterbirds is intricately linked to these environmental variables, which may vary greatly across different types of the wetlands. For example, wetland structure, such as size, shape and habitat complexity, can significantly affect waterbird distribution and behavior, with larger and more diverse wetlands often providing a greater variety of feeding, nesting and sheltering opportunities for different species [9,10]. The proximity of satellite wetlands to other natural or artificial wetlands can influence waterbird distribution by providing easier access to additional resources and potential migratory routes [11]. On the other hand, human disturbance levels, such as land development or recreational activities, can negatively impact waterbird behavior by increasing stress and disrupting nesting or feeding activities [12]. Therefore, understanding how these multi-scale environmental factors interact and affect waterbird communities in satellite wetlands is crucial when analyzing the utilization of regional wetland networks by waterbirds.
The wetlands of the middle and lower Yangtze River, particularly the Yangtze-connected lakes, are crucial wintering, breeding and stopover sites along the East Asia–Australasia Flyway, which is one of the most significant migratory routes for waterbirds [13,14]. The water level in these Yangtze-connected lakes exhibits seasonal changes, high in summer and low in winter, when large areas of mudflat and grassland are exposed. Due to the seasonal water level changes, these lakes support large numbers of migratory species, especially during the migration and wintering seasons, by providing essential feeding and resting habitats [15]. However, the wetlands (e.g., Shengjin Lake) in this floodplain have also experienced extensive degradation and loss, including aquatic vegetation decrease and water quality deterioration, mainly attributed to human activities [16]. In this context, the surrounding satellite wetlands, including artificial habitats such as aquaculture ponds, reservoirs and rice paddy fields, may play an increasingly important role in supporting these waterbird communities. These satellite wetlands may offer additional resources and refuge for waterbirds, contributing to their survival [17]. Exploring the distributions of waterbird communities in these habitats and their response to various environmental variables can provide complementary insights into how waterbirds adapt to natural wetland loss and degradation [4].
During the winter of 2022–2023 and summer of 2023, we surveyed the waterbird communities and environmental factors in the 49 satellite wetlands of four types (reservoirs, aquaculture ponds, paddy fields and natural ponds) surrounding Shengjin Lake, a Ramsar site and a typical representative of the Yangtze-connected lakes. The study aims to understand the spatial pattern of the waterbird communities and analyze the effects of environmental variables on the species’ diversity and community composition. We hypothesized that the waterbird diversity metrics (species richness, number of individuals and the Shannon–Wiener diversity index) would be higher in aquaculture ponds, and species composition would differ among various types of satellite wetlands [18]. The diversity metrics were hypothesized to be positively correlated with wetland areas, habitat diversity and landscape connectivity [9,11]. We also expected that human disturbances, calculated by distance to human settlements, would negatively affect the diversity metrics [12].

2. Materials and Methods

2.1. Study Area

Our study was carried out in the satellite wetlands surrounding Shengjin Lake, which is located in the middle and lower reaches of the Yangtze River floodplain in China (Figure 1). The climate in this area is a typical monsoonal type with an average annual temperature of 16 °C. The average annual precipitation is 1200 mm with most between May and August.
Shengjin Lake is a typical Yangtze-connected lake which experiences seasonal water level changes. The flood peaks in summer with a maximum area of approximately 1.45 × 104 ha, while the water level recedes in winter along with that of the Yangtze River. Nearly half of the lake is gradually exposed from November, providing diverse habitats, such as mudflats and grasslands, for over 70,000 waterbirds of more than 80 species each year [15]. Because of its rich biodiversity, particularly the abundant wintering waterbirds, the Shengjin Lake Nature Reserve was established in 1986 and was listed as a Ramsar site in 2015.
There are numerous satellite wetlands of different sizes and types surrounding Shengjin Lake, including reservoirs, aquaculture ponds, paddy fields and natural ponds. The reservoirs, usually small in size, are either artificially excavated or converted from natural ponds. The aquaculture ponds, ranging in size from 0.1 to 39.5 ha and surrounded by low earthen dykes, are often grouped together in continuous patches, varying in size from 2.6 to 107.8 ha. These ponds are predominantly composed of open water, with depths ranging from 80 to 150 cm during cultivation. The water is typically drained around December for harvesting, exposing the pond bottoms to direct sunlight for 30 to 60 days. The paddy fields are used for rice cultivation, with one crop harvested annually. The fields are ploughed and flooded in April, followed by rice planting in May and harvesting in October. After harvest, fallen rice grains in the unplowed fields serve as food for geese and cranes [19]. The natural ponds are less affected by human activities, maintaining more natural environmental conditions, including rich aquatic vegetation. The waterbirds recorded in Shengjin Lake are often found to use these satellite wetlands in different seasons.

2.2. Waterbird Surveys

We randomly selected 49 satellite wetlands (>1 ha) within a 4-km radius of Shengjin Lake and conducted waterbird surveys at fixed counting points on clear days during the winter of 2022–2023 and the summer of 2023. Two field surveys were carried out in each season and the time interval between surveys was approximately five weeks. During the surveys, two observers used the look–see counting method, employing binoculars (10 × 42 WB Swarovski) and a telescope (20–60 × zoom Swarovski: ATM 80) to record waterbirds at each point for 15 min. All waterbirds within the observation areas were recorded, including those flying out of the area, while birds flying in from outside the observation area were not counted [20]. No harvesting activities were observed in the aquaculture ponds or paddy fields during the survey periods. The taxonomy and nomenclature of the recorded waterbirds followed MacKinnon [21].

2.3. Environmental Variables

We measured four environmental variables for each satellite wetland (Table S1). Among these variables, wetland area (AW) and the shortest distances to main human settlements (>10 ha, DS) were calculated using high-resolution Google Earth maps. The land cover within each wetland and within its 4-km radius was interpreted from the cloud-free Sentinel-2B image (L2A-level) which was acquired on 3 January 2023 and downloaded from the ESA Copernicus Open Access Hub (https://dataspace.copernicus.eu (accessed on 8 May 2023)). We used a Support Vector Machine classification tool in ENVI 5.3 (Exelis VIS, Inc, Broomfield, Colorado, US) to recognize seven types of land cover: constructed area, mudflat, open water, aquatic vegetation, woodland, farmland and grassland. Habitat diversity (HD) was quantified by calculating the diversity of habitat types (Hill Shannon–Wiener index) within each wetland (HD; Equation (1); [22]):
H D = e − ∑ i = 1 n p i l o g p i
where pi is the area proportions of the ith type of n habitat types within each wetland. Landscape connectivity (LC) was defined as the total area of wetlands (mudflat, open water and aquatic vegetation) within the 4-km radius of each wetland. This index quantifies the wetland isolation inversely, meaning that wetlands with a greater percentage of surrounding wetlands are less isolated [23]. Except for the difference in habitat diversity within each wetland between summer and winter, other variables were the same for the two seasons.

2.4. Data Analysis

We pooled the two waterbird surveys for each wetland in each season together, respectively. Species richness, number of bird individuals and the Shannon–Wiener diversity index were calculated for each wetland in each season. Generalized linear mixed models (GLMM; [24]) with a Gaussian distribution were used to test the effects of wetland type (reservoirs, aquaculture ponds, paddy fields and natural ponds) and the environmental variables on the three metrics in each season. The two-way interaction between wetland type and each environmental factor was initially included in the models but was removed in the final models due to nonsignificant effects.
A multivariate analysis of variance (PERMANOVA) was performed using the R vegan package to test the effects of wetland types and seasons on the species’ compositions. The analysis was based on species abundance data, which were first standardized using log transformation to account for variations in sample size and species abundance. We first calculated the Bray–Curtis dissimilarity matrices to quantify the pairwise differences in species composition among the communities. The Bray–Curtis dissimilarity is a commonly used method to measure ecological distances between communities, considering both the presence and abundance of species. Given the significant effects of seasons (p = 0.001), the adonis2 function was then used to test for significant differences in community composition among the four wetland types during each season by comparing the variation explained by wetland types against the residual variation. This method utilized permutations (999 permutations in our case) to estimate the significance of the differences in community compositions. The output provided an R2 value that indicates the proportion of variation explained by wetland types, as well as a p-value to determine statistical significance. Non-metric multidimensional scaling (NMDS) was used to graphically display differences in species composition among communities in different types of satellite wetlands. NMDS used the dissimilarity matrix of species abundance to produce indirect ordination based on Bray–Curtis distances. In the NMDS diagram, communities with similar species compositions are plotted closer.
All analyses were performed using R 4.4.1, and the statistical significance level was p < 0.05.

3. Results

3.1. Waterbird Communities

We recorded 22,892 birds of 60 species in the satellite wetlands surrounding Shengjin Lake, 10,934 birds of 48 species in winter and 11,958 birds of 33 species in summer (Table 1). There were seven threatened species (two Endangered, three Vulnerable or two Near Threatened) on the IUCN Red List: oriental stork Ciconia boyciana (EN), swan goose Anser cygnoides (EN), white-naped crane Grus vipio (VU), hooded crane Grus monacha (VU), curlew sandpiper Calidris ferruginea (VU), northern lapwing Vanellus vanellus (NT) and dunlin Calidris alpina (NT). We also found nine species designated as Key Protected Wild Animal Species in China (three in the first grade: Ciconia boyciana, Grus vipio and Grus monacha and six in the second grade: Eurasian spoonbill Platalea leucorodia, common crane Grus grus, Tundra swan Cygnus columbianus, greater white-fronted goose Anser albifrons, pheasant-tailed jacana Hydrophasianus chirurgus and Anser cygnoides).

3.2. The Effects of Environmental Variables

Species richness and number of individuals differed among wetland types and were highest in aquaculture ponds in both summer and winter. The Shannon–Wiener index differed among wetland types in winter but not in summer (Figure 2).
In summer, species richness in satellite wetlands was positively affected by the wetland area, habitat diversity within wetlands and landscape connectivity. The number of individuals was positively affected by the wetland area, habitat diversity within the wetlands, and landscape connectivity, but negatively with distance to human settlements. In winter, species richness was positively affected by the wetland area and marginally by landscape connectivity. The number of individuals was positively affected by the wetland area, distance to human settlements and landscape connectivity, but negatively by habitat diversity. No effects of the measured environmental variables on the Shannon–Wiener index were detected in either season (Table 2).

3.3. Species’ Compositions

The species’ compositions differed among the waterbird communities in different types of satellite wetlands in both summer (F = 5.48, p = 0.001, R2 = 0.267) and winter (F = 3.98, p = 0.001, R2 = 0.210; Figure 3). There was also a compositional difference between each pairwise type of wetlands (Table 3). The R2 value was the highest in reservoirs and aquaculture, indicating the largest difference between these two types of wetlands in both seasons. The communities between reservoirs and natural ponds were more similar than other pairs (Table 3; Figure 3).

4. Discussion

This study highlights the critical importance of satellite wetlands around Shengjin Lake as compensatory habitats for a wide variety of waterbird species in both summer and winter. The Yangtze-connected lakes provide essential habitats for the waterbirds migrating along the East Asia–Australasia Flyway, supporting hundreds of thousands of waterbirds each year [13,25]. However, the natural wetlands have experienced extensive degradation and loss, reducing their ecological capacity for the waterbirds which have to find compensatory habitats elsewhere [13]. Due to their proximity, the surrounding satellite wetlands, which include reservoirs, aquaculture ponds, paddy fields and natural ponds, may provide essential resources such as breeding sites, roosting areas and feeding grounds for the waterbirds. Although there have been great efforts in habitat restoration inside Shengjin Lake [26], these satellite wetlands still attract many waterbirds. Our research emphasizes the role of these wetlands in maintaining the waterbird community structure and species diversity. The findings underscore the significance of preserving and managing these satellite wetlands to ensure the conservation of waterbirds and their habitats in the floodplain.
We found significant differences in waterbird communities across the four types of satellite wetlands both in diversity metrics and species composition. Reservoirs, with their relatively stable water levels and limited vegetation, supported a distinct community of waterbirds, particularly species that favor open water and deep areas for foraging [27,28]. The diversity metrics were generally the lowest in this type of wetland, particularly in winter when the water level receded in other types of wetlands but remained high in reservoirs. In contrast, aquaculture ponds, which are often managed for fish production, hosted a higher diversity of species. The pattern was more evident in winter when the bottom of the ponds was exposed, providing abundant food and a diverse range of habitats for various waterbird species [29]. The importance of aquaculture ponds as habitats for waterbirds has also been supported by other studies elsewhere [30,31,32]. Paddy fields, with their flooded rice paddies, attracted large numbers of migratory waterbirds, likely due to the abundant food supply in the form of invertebrates and seeds [33,34]. Rice paddy fields are considered as important artificial habitats for waterbirds over the world [35,36]. Along with degradation of the natural wetlands in the floodplain in the middle and lower Yangtze River, many waterbird species have been found to use rice paddy fields as foraging grounds with adjustment in behaviors and feeding habits [19,37]. Although natural ponds are closer to natural wetland ecosystems, the diversity metrics of the waterbird communities were slightly lower than those of the aquaculture ponds. This might be attributed to the fact that these natural ponds are surrounded by intense human activities [38].
The species richness and number of individuals were found to be affected by various environmental factors. Among these variables, the wetland area was positively correlated with these two metrics. This pattern is consistent with many other studies [10,39,40]. Large areas can provide more niches for more birds. In addition to wetland areas, the landscape connectivity, measured as the total area of wetlands within their 4-km radius, is also positively correlated with the two metrics. Landscape connectivity is important for wildlife in that they can easily find alternative habitats in well-connected habitat networks [41]. In the middle and lower Yangtze River, there are thousands of wetlands with large lakes (e.g., Shengjin Lake) as core habitats and satellite wetlands as compensatory habitats for waterbirds. These wetlands are connected through various ecological and hydrological processes, and those having more wetlands inside their surroundings are better connected [42]. Therefore, the well-connected satellite wetlands can support more waterbirds which can easily find alternative habitats in their surroundings. Interestingly, we found that waterbird individual numbers increased with habitat diversity and human disturbances in summer, but the pattern was in contrast to that in winter. This may be linked to the different species’ compositions between the two seasons. Gulls and egrets (e.g., little egret Egretta garzetta and whiskered tern Chlidonias hybrida) dominated the waterbird communities in summer, and these species need more diverse habitats and are usually well-tolerant of human disturbances [43]. In contrast, the waterbird communities in winter were dominated by ducks and shorebirds which need open wetlands and are vulnerable to human disturbances. The difference in habitat requirements between species in summer and those in winter also explained the positive effect of habitat diversity on species richness in summer and the nonsignificant effect in winter. Contrary to our expectation, however, we detected no effects of the measured environmental factors on the Shannon–Wiener species diversity index. This might be attributed to the fact that the effects of inter-specific competition masked the effect of environmental factors. The pattern also implies that other dimensions of diversity, such as the functional and phylogenetic diversity index, should be quantified in the future to catch the essence of biological communities [44].
Our study highlights the importance of the satellite wetlands in supporting waterbird communities, particularly in winter as a critical period when we recorded 12 threatened/protected species. Notably, we found that paddy fields could provide essential foraging habitats for Grus monachal, Anser albifrons and Antigone vipio, while aquaculture ponds provided habitats for Ciconia boyciana and Platalea leucorodia. Along with the mainstreaming of biodiversity conservation in China and the Great Protection of the Yangtze River, increased attention has been paid to the conservation of the waterbirds in the middle and lower reaches of the Yangtze River in recent decades. However, the conservation efforts are mainly directed to the large Yangtze-connected lakes with limited attention on the surrounding satellite wetlands [45,46]. We suggest that more protection efforts should be made in these wetlands, especially in aquaculture ponds, paddy fields and natural ponds, which may provide both food and shelter for the threatened species, such as Ciconia boyciana, Grus monacha, Anser albifrons and Platalea leucorodia. Meanwhile, it is important to establish a reasonable ecological compensation mechanism in the protection plans, contributing to the mitigation of potential conflicts between waterbird conservation and local communities. It should be noted that the conservation of waterbirds in these wetlands faces great challenges. Many of the satellite wetlands are not within the boundaries of any protected area and are under no conservation plans. The intense human activities, such as fishing and farming in aquaculture ponds and paddy fields, can disturb the behaviors and habitats of the waterbirds, leading to the loss of food resources and an increase in potential interferences [29]. Effective conservation requires addressing these threats through sustainable management practices where local communities should also contribute. All the wetlands across the middle and lower reaches of the Yangtze River, including natural and artificial ones, are connected together to form an integrated wetland network through various ecological and hydrological processes [47]. There are numerous satellite wetlands of various types around the natural Yangtze-connected lakes in the region which have attracted a great number of waterbirds. If the waterbirds in these satellite wetlands are threatened, the waterbird assemblages in the whole region would be in danger.

5. Conclusions

We found large numbers of waterbirds, including several threatened or nationally protected species, using the satellite wetlands surrounding the Ramsar site (Shengjin Lake) in both summer and winter. Species richness and number of individuals varied among different types of the satellite wetlands with aquaculture ponding supporting most waterbirds in both seasons. The Shannon–Wiener species diversity index exhibited similar patterns in winter but did not differ among types of wetlands in summer. Species richness was positively correlated with the wetland area, habitat diversity and landscape connectivity in summer. It was also positively correlated with the wetland area in winter. The number of individuals was positively affected by the wetland area and landscape connectivity in both seasons. It was also positively correlated with habitat diversity and negatively with distance to human settlements in summer, but the opposite pattern was found in winter. We detected no effects of environmental factors on the Shannon–Wiener species diversity. Apart from diversity metrics, species composition also differed among the waterbird assemblages in different types of the satellite wetlands in both seasons, particularly between reservoirs and aquaculture. To protect the waterbirds in the middle and lower reaches of the Yangtze River, we suggest (1) integrating the satellite wetlands into the regional wetland protection network, (2) improving landscape connectivity and enhancing habitat diversity across the satellite wetlands, (3) decreasing human disturbances, particularly in winters and (4) conducting long-term field monitoring on these waterbird assemblages and habitat variables to capture their spatial–temporal dynamics.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17030176/s1, Table S1: Numbers and the measured environmental variables of the satellite wetlands of different types surrounding Shengjin Lake; Table S2: Waterbird species recorded across the satellite wetlands of different types surrounding Shengjin Lake.

Author Contributions

Conceptualization, C.P., S.X. and G.W.; methodology, Z.Q., Q.L. and G.W.; formal analysis, C.P., Z.Q., Q.L., G.W. and T.W.; investigation, C.P., S.X. and T.W.; data curation, C.P. and G.W.; writing—original draft preparation, C.P. and G.W.; writing—review and editing, C.P., S.X., Z.Q., Q.L., G.W. and T.W.; visualization, T.W.; supervision, G.W.; funding acquisition, G.W. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Monitoring Project of Biodiversity in the Shengjin Lake National Nature Reserve (2022BFAFN02495) and the Young Elite Scientists Sponsorship Program by CAST (PhD Student Special Program, 2024).

Data Availability Statement

The data presented in this study are available on request from the corresponding author.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. The study area and the surveyed satellite wetlands.
Figure 1. The study area and the surveyed satellite wetlands.
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Figure 2. The species richness, number of individuals and the Shannon–Wiener index in the satellite wetlands of different types surrounding Shengjin Lake during summer (a–c) and winter (d–f). Pairs sharing the same subscript letter indicate no significant differences.
Figure 2. The species richness, number of individuals and the Shannon–Wiener index in the satellite wetlands of different types surrounding Shengjin Lake during summer (a–c) and winter (d–f). Pairs sharing the same subscript letter indicate no significant differences.
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Figure 3. The ordination (a, summer; b, winter) from the non-metric multidimensional scaling (NMDS) of the communities among different types of satellite wetlands surrounding Shengjin Lake.
Figure 3. The ordination (a, summer; b, winter) from the non-metric multidimensional scaling (NMDS) of the communities among different types of satellite wetlands surrounding Shengjin Lake.
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Table 1. The total species richness and number of species threatened/protected recorded in the satellite wetlands surrounding Shengjin Lake.
Table 1. The total species richness and number of species threatened/protected recorded in the satellite wetlands surrounding Shengjin Lake.
Total Species RichnessNumber of Species Threatened/Protected *
Summer
Reservoirs181/1
Aquaculture ponds271/0
Paddy fields220/1
Natural ponds210/1
Winter
Reservoirs170/0
Aquaculture ponds387/6
Paddy fields284/5
Natural ponds396/4
* The threatened species included those listed as Endangered, Vulnerable or Near Threatened on the IUCN Red List. The protected species included those designated as Grade I or II of Key Protected Wild Animal Species in China.
Table 2. The effects of the measured environmental variables on species richness, number of individuals and the Shannon–Wiener index of the waterbird communities in the satellite wetlands surrounding Shengjin Lake. The coefficient and the p-value (in the parenthesis) of each environmental variable is displayed and the significant effects are in bold.
Table 2. The effects of the measured environmental variables on species richness, number of individuals and the Shannon–Wiener index of the waterbird communities in the satellite wetlands surrounding Shengjin Lake. The coefficient and the p-value (in the parenthesis) of each environmental variable is displayed and the significant effects are in bold.
VariablesSummerWinter
Species RichnessNumber of IndividualsShannon–Wiener IndexSpecies RichnessNumber of IndividualsShannon–Wiener Index
AW0.0073 (<0.001)0.0201 (<0.001)0.0021 (0.638)0.0089 (<0.001)0.0172 (<0.001)0.0035 (0.436)
HD0.4928 (0.025)1.4540 (<0.001)0.1278 (0.810)0.1872 (0.395)−0.4728 (<0.001)0.1575 (0.775)
DS0.0001 (0.308)−0.0001 (0.0241)0.0001 (0.515)0.0001 (0.342)0.0001 (<0.001)0.0012 (0.830)
LC0.0002 (0.015)0.0005 (<0.001)0.0001 (0.735)0.0001 (0.066)0.0001 (<0.001)0.0001 (0.538)
Table 3. The results of the multivariate analysis of variance (PERMANOVA) indicating pairwise differences of species composition among different types of satellite wetlands surrounding Shengjin Lake.
Table 3. The results of the multivariate analysis of variance (PERMANOVA) indicating pairwise differences of species composition among different types of satellite wetlands surrounding Shengjin Lake.
ReservoirsAquaculture PondsPaddy FieldsNatural Ponds
Summer
Reservoirs F = 10.54, p = 0.001F = 4.45, p = 0.001F = 3.24, p = 0.003
Aquaculture pondsR2 = 0.334 F = 5.26, p = 0.001F = 6.87, p = 0.001
Paddy fieldsR2 = 0.162R2 = 0.208 F = 3.96, p = 0.001
Natural pondsR2 = 0.115R2 = 0.238R2 = 0.142
Winter
Reservoirs F = 7.11, p = 0.001F = 5.51, p = 0.001F = 2.49, p = 0.002
Aquaculture pondsR2 = 0.253 F = 4.11 p = 0.001F = 2.55, p = 0.006
Paddy fieldsR2 = 0.193R2 = 0.171 F = 3.09, p = 0.001
Natural pondsR2 = 0.091R2 = 0.104R2 = 0.114
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Pan, C.; Xu, S.; Qian, Z.; Liao, Q.; Wu, T.; Wang, G. How Do Waterbird Communities Respond to Multi-Scale Environmental Variables in the Satellite Wetlands Surrounding a Ramsar Site, Shengjin Lake in China? Diversity 2025, 17, 176. https://doi.org/10.3390/d17030176

AMA Style

Pan C, Xu S, Qian Z, Liao Q, Wu T, Wang G. How Do Waterbird Communities Respond to Multi-Scale Environmental Variables in the Satellite Wetlands Surrounding a Ramsar Site, Shengjin Lake in China? Diversity. 2025; 17(3):176. https://doi.org/10.3390/d17030176

Chicago/Turabian Style

Pan, Chengrong, Sheng Xu, Zhenbing Qian, Qichen Liao, Tongxinyu Wu, and Guangyao Wang. 2025. "How Do Waterbird Communities Respond to Multi-Scale Environmental Variables in the Satellite Wetlands Surrounding a Ramsar Site, Shengjin Lake in China?" Diversity 17, no. 3: 176. https://doi.org/10.3390/d17030176

APA Style

Pan, C., Xu, S., Qian, Z., Liao, Q., Wu, T., & Wang, G. (2025). How Do Waterbird Communities Respond to Multi-Scale Environmental Variables in the Satellite Wetlands Surrounding a Ramsar Site, Shengjin Lake in China? Diversity, 17(3), 176. https://doi.org/10.3390/d17030176

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