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Response of chlorophytes to environmental variables in field vs. forest pondsOur study demonstrated that human-originated transformation in the catchment area surrounding a small water body may influence the water conditions in terms of physical, chemical, and biological parameters as well as the ecological state of the aquatic environment in respect to green algae communities.Chlorophytes inhabiting field ponds were more abundant compared with the forest ponds. This shows that field ponds, due to the higher values of TRP and water conductivity, created favorable conditions for chlorophyte development. The high concentrations of TRP and conductivity in aquatic environments are characteristic in the case of agricultural catchments exposed to anthropogenic pressure because of the inflow from the surrounding fertilized fields42. In this type of pond, we also observed significantly higher water temperatures and pH due to the lack of trees around them compared to the forest ponds, two factors which also positively influenced the growth of chlorophytes. Both the higher light intensity and the smaller size of the field ponds cause earlier warming up than the forest ponds and give an advantage to high light tolerant species. Moreover, it is well known that an increase in temperature stimulates the release of phosphorus from the bottom sediments, so this could be another reason for the higher levels of TRP in the field ponds. Our CCA analysis showed that TRP and conductivity were the strongest determinants of the distribution of chlorophyte species in the examined water bodies. We found a large group of dominant species indicated high values of TRP (e.g. Ankistrodesmus falcatus, A. arcuatus, Monoraphidium griffithii, Pseudopediastrum boryanum, Pediastrum duplex, Scenedesmus obtusus, Scenedesmus arcuatus var. gracilis, Desmodesmus communis, Coelastrum microporum), and another group of species (e.g. Kirchneriella irregularis var. spiralis, Tetraedron minimum, Scenedesmus ecornis) that preferred high levels of conductivity.In the field ponds generally higher mean abundances of filtrators and Rotifera were observed. This could be another important factor stimulating the growth of chlorophytes and increasing their abundances by the resupply of nutrients through excretion43,44. On the other hand, the high densities of algae could be the factor that caused better zooplankton development, and therefore its abundance in field ponds was greater. Filtrating cladocerans and Rotifera also had a significant influence on the distribution of chlorophyte dominating species. However, even though the total abundance of both chlorophytes and filtering zooplankton was greater in the field ponds, CCA analysis revealed a negative relationship existing between filtrators and most dominant species of chlorophytes (e.g. Pandorina morum, Willea rectangularis, Desmodesmus armatus, Nephrochlamys willeana, Cosmarium trilobulatum). Only two chlorophyte species—Lemmermannnia tetrapedia and Tetraedron triangulare—co-occurred with cladoceran zooplankton. These latter species are very small compared to the species above and can therefore be overlooked by filtrators, which have a choice of larger and perhaps more nutritiously satisfying algae of the genus Pandorina, Crucigeniella, Cosmarium or Nephrochlamys, but still of a size suitable for zooplankton. It can also be interpreted in such a way that Crucigenia and Tetraedron are among the r-strategists that reproduce very quickly, so grazing pressure by zooplankton can stimulate their rapid development45 and thus they remain at a stable level.Specific environmental conditions prevailing in the field ponds resulted in a high number of exclusive taxa44, found only in this type of water body. Moreover, a greater diversity of the representatives of different functional groups were found here, compared to the forest ponds.Analyzing the distribution of chlorophytes in terms of phytoplankton functional groups39,40, we found that group W1 was represented by only one species, Gonium pectorale. This was especially noted in the field water bodies. This group is known to prefer small water bodies rich in organic matter from husbandry or sewage40, which suggests that the field catchment in our study migh be a supplier of these substances. It also proves that field surroundings are far more human impacted. In the field ponds we observed a higher abundance of chlorophytes belonging to the groups G (Eudorina elegans, Pandorina morum, Pandorina smithii and Volvox aureus), J (e.g. representatives of the genus Actinastrum, Chlorotetraedron, Coelastrum, Crucigenia, Desmodesmus/Scenedesmus, Golenkinia, Pediastrum, Tetraedron, Tetrastrum, Westella, Willea/Crucigeniella), W0 (genera Chlamydomonas, Chlorangiopsis, Chlamydomonadopsis, Planktococcomyxa/Coccomyxa) and X3 (Chlorella sp.), typical for shallow nutrient-rich waters (G and J), ponds with extremely high organic contents (W0), and for shallow well-mixed layers (X3), according to classification given by Padisak et al.40. Considering that nitrogen compounds had a similar level in both types of ponds it can be stated that the representatives of the above mentioned functional groups of chlorophytes associated with the field ponds were presumably dependent on higher concentrations of TRP and conductivity and not that much on nitrogen concentrations.In the forest ponds significantly higher values of water saturation were recorded compared to the field ponds. Moreover, the lack of inflow of fertilizers from the catchment area resulted in lower TRP concentrations, which along with lower water temperatures, pH and conductivity in the forest ponds may have contributed to the reduced abundance of chlorophytes compared to the field water bodies. RDA analysis showed that some dominant chlorophyte species (e.g. Closterium moniliferum, Closterium tumidulum, Cosmarium trilobulatum and Mougeotia sp.) were associated with this type of small water body. At the same time the abundance of these species was smaller in the field ponds. We also found that chlorophyte diversity (Shannon–Weaver index) was greater in the forest ponds. This suggests that water bodies located within the forested area, usually more natural ponds being less exposed to anthropogenic pressure, are characterized by greater biodiversity. Moreover, in this type of water body we found many exclusive species39, not reported from the field ponds. Interestingly, about the half of these taxa belonged to desmids, which prefer lower pH and conductivity46, conditions typical for forest ponds. This could be also a reason for the dominance of desmid species with the highest abundance/frequency, associated with forest ponds.Taking into consideration the phytoplankton functional groups39,40 our study showed that the chlorophytes associated with forest ponds prefer mesotrophic waters (from the group TD: Cladophora glomerata, Geminella turfosa, Geminella planctonica, Microspora sp., Netrium digitus, Oedogonium sp., Oocystidium ovale, Spirogyra sp. Zygnema sp. and those belonging to the group N: mainly genera Closterium, Cosmarium, Euastrum, Micrasterias, Staurastrum, Staurodesmus, Xanthidium). This explains their greater share in the less fertile forest ponds. Another group associated with the forest ponds – T (Mougeotia sp., Binuclearia lauterbornii) contains species tolerant to light deficiency, so they were able to develop well in the more shaded water bodies located in the forest catchment.Chlorophyte community structure in two types of habitats (open water vs. macrophyte-dominated zone)In our study, the type of habitat (open water and macrophyte-dominated zones) also had a significant structuring effect on chlorophytes. There were a group of species linked to the open water zone (Pandorina morum, Nephrochlamys willeana, Oocystis lacustris, Scenedesmus armatus, Scenedesmus intermedius and Desmodesmus communis), being negatively related to vegetated stations at the same time. Generally, we found here a higher mean abundance of chlorophytes compared to the macrophyte-dominated zones, possibly due to the higher values of nutrients such as NH4 and TRP, the conditions favouring the development of many algae species. The results of the CCA analysis with habitats confirmed the high importance of both nutritional factors in structuring the distribution of chlorophyte species. There was a group of species associated with a rise in the concentration of ammonium (e.g. Scenedesmus arcuatus var. gracilis, Pediastrum duplex, Closterium moniliferum, Closterium tumidulum, Cosmarium trilobulatum, Willea rectangularis) as well as with phosphates (Monoraphidium tortile, Scenedesmus ecornis, Tetradesmus lagerheimii and Tetraedron minimum). Generally, high abundance of chlorophytes in the open water area was accompanied by a small-sized fraction of zooplankton–rotifers. Therefore, rotifers had a lower impact on the distribution of chlorophytes than filtrators. The increasing numbers of cladocerans contributed to the lowering abundance of some chlorophytes, such as Monoraphidium tortile, Scenedesmus ecornis, Tetradesmus lagerheimii or Tetraedron minimum. This shows that filtrators, whose densities were significantly higher among macrophytes, were able to control the development of some chlorophyte species much more efficiently than small-bodied rotifers.The effect of habitat was also visible in the case of phytoplankton functional groups39,40. We found that representatives of the group N (e.g. Closterium, Cosmarium, Euastrum, Micrasterias, Staurastrum) had a significantly higher mean abundance in the open water zones compared to the macrophyte-dominated zones. Interestingly, according to Padisak et al.40 group N prefers less fertile (mesotrophic) conditions, which is inconsistent with our results. However, we think that their association with the open water sites could be connected rather with the place/level where they live in the water column, rather than with the trophic state of water. The above mentioned chlorophytes taxonomically belong to desmids, which are mostly benthic organisms. Their greater quantitative share in the samples from the open water areas could be an effect of the intensive water mixing in the shallow ponds due to the lack of macrophytes. Neustupa et al.47 confirm that desmids are able to form tychoplanktonic communities due to water movements. In the samples collected from the macrophyte-dominated stations the mean abundance of desmids was generally lower, probably because of the macrophyte stabilizing effect. Aquatic plants are known to reduce turbidity and stabilize bottom sediments48, so they can prevent any intensive water mixing in ponds. In the examined open water stations, we also found a higher mean abundance of chlorophytes typical for shallow nutrient-rich waters (group G: Eudorina, Pandorina, Volvox and group K: Radiococcus) and/or for ponds with extremely high organic contents (group W0: e.g. Chlamydomonas), which proves that the sites lacking macrophytes were more fertile. Additionally, clearly more representatives from the codon J and X1 (typical for waters with high trophic levels) and a greater diversity of the representatives of different functional groups were recorded in the open water area compared to the macrophyte-dominated zones.The macrophyte-dominated stations had more abundant communities of filtrators, as aquatic plants are known to provide a profitable shelter for zooplankton49. Cladoceran predominance among macrophytes may have been a force reducing green algae numbers. The chlorophytes of the investigated ponds were mostly small- or medium-size species. Their size distribution makes them a high quality food for zooplankton, particularly for cladoceran filtrators. According to RDA analysis apart from pond size, the presence of filtrators significanly reduced the abundance of several chlorophyte dominating species. The lower algae abundance among macrophytes compared to the open water zone could also be explained by competition between algae and macrophytes for light and nutrients37,50 and/or with the secretion of allelopathic substances e.g. by Ceratophyllum demersum51 inhibiting algal development. Our studies demonstrated that among chemical factors which clearly differentiated the two types of analysed habitat, TRP and NH4 significantly influenced the distribution of chlorophyte dominating species. The lower levels of these parameters in macrophyte-dominated zones suggest that the nutrient uptake by aquatic plants in the investigated water bodies was high. There are many reports on the decrease of nutrient concentrations by macrophytes30,37,52, which are consistent with our observations. Despite lower, compared to the open water zone, chlorophyte densities within the macrophyte-dominated zones there was a group of species (e.g. Mougeotia sp., Pediastrum tetras, Scenedesmus obtusus, Monoraphidium contortum) that selectively chose vegetated stands. Furthermore, we found a great number29 of exclusive chlorophyte species for macrophyte-dominated zones. Half of these taxa belong to desmids, which are often periphytic organisms associated with aquatic macrophytes53,54.Preference towards macrophyte-dominated stations was also documented for two phytoplankton functional groups (T: Mougeotia sp. and Binuclearia lauterbornii and TD: e.g., Spirogyra sp., Zygnema sp., Cladophora glomerata, Oedogonium sp.) and one group which occurred exlusively among vegetated sites (MP—Ulothrix). Interestingly, all the representatives of these groups had a similar filamentous morphological form, which suggests that many of them are of epithytic origin, coexisting within aquatic plants. Two more groups—X2 (Pseudodidymocystis/Didymocystis, Pteromonas) and W1 (Gonium pectorale) were clearly affected by the presence of macrophytes. According to Padisak et al.40, codons TD and X2 indicate mesoeutrophic conditions and their higher abundances in the macrophyte-dominated zones also proves that plants contribute to lowering the trophic levels in the examined ponds. On the other hand, the relatively high abundance of the representative of the group W1 in these habitats suggests that macrophytes could enrich ponds with organic matter during the process of their decomposition.Concluding, our results prove that different types of catchment area (field and forest) as well as different types of habitats (open water zone and macrophyte-dominated zone) create distinct, specific conditions (dependent on some physical–chemical and biological variables) for the occurrence of chlorophytes in small water bodies. We conclude that cosmopolitan chlorophytes undoubtedly respond to the level of habitat heterogeneity, contributing to the ecological assessment of small water bodies. Chlorophytes in particularl react to the level of human transformation in the ponds’ vicinities. This is why we suggest using them for water quality evaluation in ponds. This interdisciplinary research significantly broadens the knowledge, not only about the response of chlorophytes to physical–chemical parameters of water, but also about the food preferences of zooplankton for which green algae are the basic food, and vice versa about the impact of zooplankton on microalgae communities. The analyses provide valuable information on chlorophytes-zooplankton interactions and also about the relationships between chlorophytes and macrophytes. Received data emphasize the high value of field ponds, underestimated habitats particularly vulnerable to destruction in the agricultural landscape. The research will help to better understand the functioning of poorly studied small water bodies, which will contribute to the preservation of their biodiversity and protection against degradation. They will also be useful in the management of small water bodies based on the specificity of chlorophyte occurrence in various habitats and catchment type ponds. Moreover, these results are important in a broader context, as the interactions between the studied organisms and the physico-chemical parameters of water in small bodies of water are to some extent universal, so the analyses will broaden the knowledge about the functioning of larger bodies of water. More

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Mixing fire occurrence with wildfire activity is problematic also when trying to draw policy conclusions. Fang et al.1 examined the temporal pattern of fire numbers between 2005-18 and concluded that the application of a fire suppression policy after 1987 has contributed to decreases in fire occurrences after 2007. However, fire suppression is an effort to mitigate the results of a fire once it has started10. Consequently, fire suppression strictly affects the burned area, and not fire occurrence. Other aspects associated with fire planning, like awareness campaigns or fire bans, may act on fire occurrence. However, any relationship between fire occurrence and fire suppression will necessarily be artefactual because the latter does not affect the former.We acknowledge that part of the discrepancy with Fang et al.1 may lie in the different scales used in these analyses. However, fire activity is a term that currently lacks a rigorous definition and should be used with caution. Fire occurrence depends primarily on the number of ignitions (along with other factors affecting fire detection such as climate, topography or vegetation), which, in turn, results from human activity1 and, in some areas, lightning11. Using fire occurrence as an indicator for fire activity is particularly problematic when comparing multiple biomes that show marked differences in fire regime, as we demonstrate here. Additionally, ENSO and fire suppression may both affect burned area, but there is currently no mechanism that can explain a mechanistic link between either of these processes and the number of fire events. Consequently, fire occurrence should not be used as a sole metric of fire activity.We additionally note that burned area is not necessarily a reliable metric of fire impacts on ecosystems and society. Significant variation in severity and intensity may occur within a fire perimeter12. Additionally, damage to people and property are not captured by this metric13. While we caution against the use of a single metric to evaluate fire activity, we hope to have demonstrated that using fire occurrence alone is particularly problematic, and that the picture it paints is rather unrealistic. More

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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