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Species composition of vegetation in the WLFZIn this survey, a total of 44 species in 43 genera of 21 families of vascular plants were found and confirmed in the reservoir WLFZ of the Jinsha River basin, among which, 13 genera and 13 species of Compositae, 4 genera and 4 species of Gramineae, 3 genera and 3 species of Amaranthaceae, 2 genera and 2 species of Verbenaceae, Labiatae, Umbelliferae, Cruciferae and Convolvulaceae, 1 genus and 2 species of Polygonaceae, and the remaining 12 families were all single genera. Compositae had the highest number of species, followed by Gramineae and Amaranthaceae, accounting for 29.55%, 9.09% and 6.82% of the total number of species in this survey, respectively, which are the main dominant families in the region.According to the life type classification system of the Flora of China, the plants in the WLFZ of this survey can be classified into five life types: annual herbs, perennial herbs, annual or biennial herbs, annual or perennial herbs, and biennial herbs. The community is overwhelmingly dominated by annuals with a high proportion of 54.55%, followed by perennials with 34.09% and the rest of all life types with a total of 11.36%.The higher number of annual plants indicates that the environmental conditions in the WLFZ are harsher after inundation by water storage, and plants that can complete their entire life cycle in a short period of time after receding water are more likely to survive compared to plants that take a long time to complete their entire life cycle.The vegetation types in each study area of the WLFZ are shown in Table 3, among which 17 species, including S. subulatum, E. humifusa, C. bonariensis, V. officinalis, O. biennis, S. plebeia, U. fissa, B. juncea, S. orientalis, D. repens, A. lividus, T. mongolicum, G. parviflora, P. praeruptorum, P. hys-terophorus, D. stramonium and Ph. Nil, are newly discovered species in the reservoir WLFZ, which are rarely reported in other reservoir WLFZ studies so far. Among the study areas, the Longkou study area was the richest in vegetation types, with the most families, species and life types among all study areas, and the number of perennial herb species was comparable to that of annual herb species, while all other study areas were mainly dominated by annual herbs. The vegetation composition of the remaining study areas averaged 6–8 families and 11–12 species, except for the Ludila study area with no plants growing and the Liyuan study area with only 5 families and 5 species. In general, each study area was dominated by Compositae and Gramineae.Table 3 Vegetation composition in each study area.Full size tableVegetation area, coverage, and percentage of the WLFZAccording to the vegetation classification in the WLFZ of each study area (Fig. 5 and Table 4), the vegetation coverage of the study areas of the Liyuan, Ahai, Ludila and Guanyinyan reservoir WLFZ were all less than 5%. The study area of Ludila was completey devoid of vegetation in the WLFZ. The coverage in Liyuan was only 0.02%, with mostly individual herbaceous plants sporadically distributed on the upper boundary of the WLFZ. In Ahai, C. dactylon grow concentratly in patches at the top of the WLFZ together with some other sparsely growing vegetation, with a coverage of 1.47%. The vegetation coverage of Guanyinyan was 3.21%, mainly distributed in the upper part of the WLFZ and expanding towards the middle. In this area, 30.39% of the vegetation was X. sibiricum, growing in large tracts as low seedlings; 21.03% was A. sessilis growing in patches, 10.87% was C. dactylon growing mainly on the upper boundary of the WLFZ, and 37.71% was a mixture of plants growing in clusters with only a few of each.Figure 5The results of vegetation classification in the WLFZ of each study area. (a) Liyuan, (b) Ahai (c) Longkaikou, (d) Ludila, (e) Guanyinyan, (f) Xiluodu. Note: Non-Veg (Non-vegetation), Other-Veg (Other vegetation), C. Dac (Cynodon dactylon), A. Ses (Alternanthera sessilis), C. Bon (Conyza bonariensis), Ch. Amb (Chenopodium ambrosioides), C. Can (Conyza canadensis), D. Rep (Dichondra repens), H. Sib (Hydrocotyle sibthorpioides), V. Off (Verbena officinalis), X. Sib (Xanthium sibiricum). (Generated with eCognition Developer, and the URL is https://www.ecognition.com).Full size imageTable 4 Vegetation area, vegetation coverage and vegetation classification accuracy of WLFZ in each study area.Full size tableThe vegetation coverage of Longkaikou and Xiluodu WLFZ was more abundant, 46.47% and 55.81% respectively. In Longkaikou, vegetation mainly covered the middle and upper parts of the WLFZ. Of the vegetation, 66.38% was C. dactylon, 26.50% was A. sessilis, 2.35% was H. sibthorpioides, 1.68% was Ch. ambrosioides, and 3.09% was a variety of vegetation species, only a few of each, divided into Other-Veg class.Due to weather and equipment constraints, we were unable to photograph the upper and lower boundaries of the WLFZ in Xiluodu study area, but we still obtained the images of the main part of the WLFZ, which consisted mainly of 58.4% X. sibiricum, 28.04% C. dactylon, 10.59% S. viridis, and 2.97% other vegetation.The vegetation coverage in the WLFZ of different reservoirs of the Jinsha River basin varied significantly, but in terms of quantity, most of them were absolutely dominated by 1–4 species, which were distributed in patches and strips, and covered an area and proportion far more than the rest of the vegetation, while the rest of the vegetation was sparse in quantity each and was sporadically distributed. C. dactylon, A. sessilis, X. sibiricum, S. viridis, H. sibthorpioides, Ch. Ambrosioides were the main dominant and pioneer species for vegetation restoration in the reservoir WLFZ of the Jinsha River basin.Spatial distribution pattern of vegetation in fluctuating zoneSince no vegetation survived in the Ludila study area, and the vegetation in the Liyuan, Ahai and Guanyinyan study areas was sparse, with less than 5% coverage, and all of them were concentrated in the upper part of the WLFZs (Fig. 5), this paper mainly analyzed the spatial distribution pattern of vegetation in the Longkou and Xiluodu study areas, which had better vegetation coverage.Landscape patternCA is a basic index for landscape pattern study, and LPI reflects the proportion of the largest patch in the landscape type to the total landscape area, which is an expression of patch dominance. The SHAPE and PAFRAC describe the complexity of patch shape, the larger the SHAPE value indicates the more complex patch shape; the closer the PAFRAC value to 1 indicates the more regular patch shape. PROX reflects the degree of proximity of each landscape type, the larger its value indicates the higher degree of patch aggregation and the lower degree of fragmentation; ENN describes the degree of physical connection of the landscape types, the larger its value indicates the greater distance between patches and the greater degree of fragmentation.From the overall landscape level (Fig. 6), in the Longkaikou study area, CA and LPI showed that the areas of vegetation patches were large, less spatially fluctuating and uniform distribution, with obvious patch dominance, reflecting characteristics of patchy distribution; PROX and ENN showed that the vegetation patches were clustered and the landscape was well connected; SHAPE and PAFRAC showed that there was little variation in the shape complexity of vegetation patches in most areas of the WLFZ.Figure 6Spatial characteristics of vegetation landscape pattern index in the Longkaikou study area (Generated with ArcGIS 10.5 software, and the URL is: https://www.esri.com/en-us/home).Full size imageAt the level of landscape types (Table 5), the vegetation landscape types in the Longkou study area included C. dactylon, A. sessilis, H. sibthorpioides and other vegetation, among which, C. dactylon showed significant advantages in patch area, patch dominance, patch aggregation and connectivity; followed by A. sessilis and H. sibthorpioides, A. sessilis was significantly better than H. sibthorpioides in patch area, but in patch shape, H. sibthorpioides was more aggregated than A. sessilis and had better patch connectivity; Other-Veg showed significant weaknesses in patch area and aggregation; there were no significant differences among the landscape types in patch shape.Table 5 Landscape index of patch types in the Longkaikou study area.Full size tableThe spatial characteristics of the vegetation landscape pattern index in the Xiluodu study area were shown in Fig. 7. From the overall level of the landscape, the area of vegetation patches and the dominance of patches were spatially variable, the vegetation was well connected, with obvious characteristics of patchy distribution, and the shape of vegetation patches did not show obvious spatial characteristics.Figure 7Spatial characteristics of vegetation landscape pattern index in the Xiluodu study area (Generated with ArcGIS 10.5 software, and the URL is:https://www.esri.com/en-us/home).Full size imageFrom the level of landscape types (Table 6), the vegetation landscape types in Xiluodu study area included four categories: X. sibiricum, C. dactylon, S. viridis and Other-Veg type. Among them, X. sibiricum showed obvious advantages in patch area, patch dominance, patch aggregation and connectivity, followed by C. dactylon, both of which were significantly better than S. viridis and Other-Veg, and the differences in patch shape complexity among landscape types were small.Table 6 Landscape index of patch types in the Xiluodu study area.Full size tableDistribution characteristics along terrainAccording to the statistics (Fig. 8), the vegetation area share of Longkaikou study area in the upper, middle and lower elevation gradients of the WLFZ was 54.61%, 26.62% and 18.77%, respectively, indicating that the vegetation was mostly in the upper part of the WLFZ, with a coverage of 83.80%, while the vegetation in the lower part was the least, with a coverage of less than 1%. From the viewpoint of each vegetation species, in the upper part of the WLFZ, C. dactylon had the largest area, accounting for 66.9% of the total vegetation area, followed by A. sessilis, accounting for 25.9%, while H. sibthorpioides and Other-Veg only survived in the upper part, accounting for 2.3% and 4.9% each. From the distribution of each slope class, the vegetation of the WLFZ gradually decreased with the increase of slope, and the vegetation was mainly concentrated in the range of slope 35°, and the coverage of each vegetation decreased significantly when the slope exceeded 35°. In the aspect, the distribution of vegetation in the WLFZ did not show any obvious preference. The surface relief in the study area of Longkou was generally low, and C. dactylon was mainly distributed in the range of surface relief less than 0.84 m. When the surface relief is greater than 2.52 m, the vegetation coverage tends to be close to 0. The vegetation showed no obvious distribution preference in terms of surface roughness and topographic wetness index.Figure 8Changes in vegetation coverage with topographic factors in the Longkaikou study area (Drawn with Origin 2018_64Bit, and the URL is https://www.OriginLab.cn/).Full size imageThe spatial distribution of vegetation in the study area of Xiluodu was shown in Fig. 9. The maximum drop in water level at Xiluodu study area can reach 60 m, but only the half of the upper part of the subsidence zone with a drop of about 30 m was photographed. The coverage rate of C. dactylon was the largest in this elevation gradient, S. viridis was mainly distributed in the uppermost part of the zone, while X. strumarium was well covered in all elevation gradients. From the distribution of surface relief, the overall vegetation coverage decreases with the increase of surface relief, with X. strumarium and S. viridis mainly distributed in the area of 0–3.45 m, while both the coverage of C. dactylon and Other-Veg were not much different across the surface relief . The distribution of vegetation showed no obvious preference in terms of slope, aspect, surface roughness and topographic wetness index.Figure 9Changes in vegetation coverage with topographic factors in the Xiluodu study area (Drawn with Origin 2018_64Bit, and the URL is https://www.OriginLab.cn/).Full size imageInfluence of topographic factors on the spatial distribution pattern of vegetation in the WLFZAccording to the results of species distribution modeling, the number of samples in the study area of Longkaikou was 39,321, and the overall accuracy of the model was 88.2%. The terrain factors, in descending order of importance, were elevation  > slope  > surface relief  > surface roughness  > aspect  > topographic wetness index, with values of 0.681, 0.146, 0.091, 0.042, 0.033 and 0.007, respectively (Fig. 10). It can be seen that the vegetation distribution in the WLFZ was mainly influenced by elevation, followed by slope and surface relief, and is less influenced by surface roughness, aspect and topographic wetness index. This was consistent with the results of typical correlation analysis.Figure 10Ranking of important values of topographic factors in the Longkaikou study area (Drawn with Origin 2018_64Bit, and the URL is https://www.OriginLab.cn/).Full size imageA total of six pairs of typical variables were calculated in the Longkou study area, and standardized typical coefficients were used due to the inconsistency of each landscape pattern index as well as topographic factor units. According to the results of significance test (Table 7), the first four pairs of typical p-values were less than 0.05, indicating that the correlations reached a significant level, and their correlation coefficients were 0.565, 0.262, 0.142, and 0.034, among which the correlation coefficient of the first pair was the largest, so the first pair was selected for analysis. The topographic factors and landscape indices highly correlated with the first pair of typical variables were elevation, surface relief and CA and SHAPE, respectively. According to Tables 8 and 9, their mechanism of action was that the greater the elevation, the smaller the surface relief, resulting in a larger patch size and more complex shape of the vegetation, and therefore a more frequent exchange of energy with the outside world and a greater ability to survive.Table 7 Significance test of typical correlation coefficient in the Longkaikou study area.Full size tableTable 8 Standardized canonical correlation coefficients of terrain factors in the Longkaikou study area.Full size tableTable 9 Standardized typical correlation coefficients of landscape pattern in the Longkaikou study area.Full size tableThe number of samples in the study area of Xiluodu was 41,010, and the overall accuracy of the model was 61.4%. The terrain factors, in descending order of importance, were elevation  > surface relief  > ground roughness  > aspect  > slope  > terrain moisture index, with values of 0.395, 0.209, 0.157, 0.123, 0.073, and 0.043, respectively (Fig. 11). It can be seen that the vegetation distribution in the WLFZ was most influenced by the elevation, followed by the surface relief.According to the typical correlation analysis, six pairs of typical variables were calculated for the Xiluodu study area, of which the first four pairs had typical P values less than 0.05 (Table 10), indicating that the correlation reached a significant level, and their correlation coefficients were 0.299, 0.208, 0.102, and 0.033, and the first pair was the largest, so the first pair was selected for analysis.The topographic factors and landscape indices with high correlation with the first pair of typical variables were elevation,surface relief and CA, PAFRAC, respectively, and according to Tables 11 and 12, their mechanism of action was that the greater the elevation, the greater the surface relief, leading to a smaller patch area and simpler shape of the vegetation.Figure 11Ranking of important values of topographic factors in the Xiluodu study area (Drawn with Origin 2018_64Bit, and the URL is https://www.OriginLab.cn/).Full size imageTable 10 Significance test of typical correlation coefficient in the Xiluodu study area.Full size tableTable 11 Standardized canonical correlation coefficients of terrain factors in the Xiluodu study area.Full size tableTable 12 Standardized typical correlation coefficients of landscape pattern in the Xiluodu study area.Full size tableLimiting factors of vegetation restoration in WLFZPreliminary studies showed that after long-term water level fluctuations in the cascade reservoirs, most of the vegetation in the WLFZs of the cascade reservoirs in the Jinsha River basin could be restored to different degrees, however, the restored species types were relatively simple, all of them were herbaceous plants, and mainly annual herbaceous plants. The restoration of the WLFZs of different reservoirs varied significantly, with vegetation coverage of more than 46% and 27 species types in the better restored areas, such as the Longkou study area, while the vegetation coverage of the less restored areas was usually less than 5% and 5–12 species types, and some areas even had no grass, such as the Ludila study area. According to the statistics (Fig. 12), the habitats in the study area of different reservoirs in the Jinsha River basin were significantly heterogeneous, with significant differences in climate, soil conditions, topography, and water level drop, etc. Because of the inconsistent range of values and units of different environmental factors, comparative analysis was performed by normalization, as shown in Fig. 12, vegetation cover was significantly correlated with the average soil Ph and the average thickness of the subsurface 30 cm soil layer, and the two study areas with average soil Ph greater than 8, Pear Garden and Rudyra, were almost completely bare. These two study areas were almost dominated by sand and gravel, with thin soils averaging  8 and soil thickness  More

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