Changes and driving factors of psychrophilic diatom community structure of algal mats in Huanglong Scenic Area
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摘要: 硅藻群落对环境变化十分敏感，是自然河流水质变化的指示生物。为了解黄龙风景区藻席及藻席嗜冷硅藻群落结构与环境变量的关系，对景区内8个典型景点进行水样和藻席的采集并分析。结果表明，黄龙风景区藻席是以钙华作为基底，丝状藻作为骨架，栖息着大量硅藻的高寒岩溶微生态系统，共鉴定出25属72种硅藻，其中优势种13种，以桥弯藻属（Cymbella）为主。通过冗余分析发现，水温（F = 2.8，P = 0.02）和溶解硅（F = 2.3，P = 0.04）是影响黄龙风景区藻席嗜冷硅藻群落结构特征的主要驱动因子，TN是潜在驱动因子。硅藻可以为高寒岩溶区地表水监测提供重要的补充信息，建立完整的硅藻物种数据库，有利于区域水质监测。Abstract: Huanglong Scenic Area, located in Songpan county, Sichuan Province, China, is listed as the World Natural Heritage, and is covered with travertine landscape at an altitude of about 3,145-3,588 m. With the plateau temperate sub-frigid monsoon climate, the melt water from snow mountains, the atmospheric precipitation and the underground karst water are the main water sources of Huanglong Scenic Area. In addition, covered by ice and snow at extremely low temperatures, Huanglong Scenic Area has a half-year-long freezing period with an annual average temperature of only 1.1℃. A large number of algae, mainly psychrophilic diatoms, grow in the water coverage of the travertine deposition area in the scenic area. The diatom community, an indicator of water quality change, is very sensitive to environmental changes in natural rivers. At present, there are still few studies related to this alpine karst area. In order to explore the structure of algal mats as well as the relationship between the community structure of psychrophilic diatoms in algal mats and environmental variables in Huanglong Scenic Area, we analyzed water samples and algae mats from 8 typical scenic spots in October 2021. The water temperature (T), pH and conductivity (Ec) as well as the concentrations of ${\rm{HCO}}_3^{-}$ and ${\rm{CO}}_3^{2-}$ of the sampling points were measured on site. At the same time, the geographic data of the sampling points were recorded. The concentrations of dissolved silicon (Dsi), cations (Ca2+, Mg2+, Al3+, K+), anions (Cl−, ${\rm{NO}}_3^{-}$, ${\rm{SO}}_4^{2-}$), TN and TP were determined in the laboratory. The structural compositions of the algal mats and the characteristics of the diatom community structure were analyzed. The main driving factors affecting the water of Huanglong Scenic Area were judged by Principal Component Analysis (PCA), and one of the two environmental variables with greater correlation was eliminated by Spearman Correlation Analysis. After Detrend Correspondence Analysis (DCA) of diatom species abundance in algal mats, Redundancy Analysis (RDA) was selected to analyze the relationship between environmental variables and diatom communities. The results show that the concentrations of Ca2+ and ${\rm{HCO}}_3^{-}$ in the waterbody is high, and water mainly belongs to the HCO3-Ca type, followed by the HCO3-Ca-Mg type. The algal mats are mainly composed of travertine particles, filamentous algae and diatoms. The study area is an alpine karst micro-ecosystem with travertine as the base and filamentous algae as the skeleton, inhabiting a large number of diatoms. 72 species of diatoms from 25 genera are identified from algal mats, among which 16 species have a total relative abundance greater than 2%. The genus Cymbella (14 species), Diatoma. (8 species), Cymbopleura (7 species), Caloneis (4 species) and Synedra (4 species) contribute the most. There are 13 dominant species of diatoms in Huanglong Scenic Area. 4 species from the genus Cymbella have absolute advantage (Ytotal = 0.132) at the genus level. At the species level, there is little difference in the dominance index of each dominant species (0.023≤Y≤0.053), among which the dominance index of Diatoma vulgaris is the largest, and that of Cymbopleura linearis is the smallest. In terms of the diversity index, there are some differences in the Shannon-Weiner index, Simpson diversity index and Pielou index at various sampling points. As the altitude decreases, the diversity index from #1 to #8 fluctuates, rising and then falling repeatedly. This may be related to the water circulation system of Huanglong Scenic Area. The analyses of the relationship between environmental variables and diatom communities by RDA show that the eigenvalues of the first two axes are 0.43 and 0.24, respectively, explaining 67.68% of the cumulative variance of data about diatom community species. The correlation between environmental variables and diatom community structure in algal mats is T>DSi>pCO2>TN> Mg2+>Ca2+>altitude, and the three environmental variables of Cl−, ${\rm{NO}}_3^{-}$ and ${\rm{SO}}_4^{2-}$ are excluded because of the low correlations. The significance test of the influence degree of each factor by Monte Carlo test finds that T (F=2.8, P=0.02) and DSi (F=2.3, P=0.04) are environmental variables that significantly affect the algae community structure of the algal mats. In addition, although TN is not the most significant driving factor affecting diatoms of the algal mats in Huanglong Scenic Area, there is a trend of eutrophication in the waterbody of the scenic spot, which will affect the algal community structure, so TN may be a potential driving factor. The above conclusions can provide a basis for the establishment of the diatom species bank in Huanglong Scenic Area and the monitoring and management of the river water environment in the plateau karst area.
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Key words:
- Huanglong Scenic Area /
- algal mats /
- diatoms /
- community structure /
- driving factors /
- environmental variables
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图 1 黄龙风景区位置及采样点分布
Figure 1. Distribution of sampling points in Huanglong Scenic and Historic Interest Area
图 2 藻席结构和组成解析
a-b. 使用SEM-EDS分析藻席微观结构; c. 使用元素分析仪分析不同颜色藻席的C，H，N，S元素含量; d. 使用FTIR分析不同颜色藻席的官能团或化学键; e. 使用XRD分析藻席中的矿相成分。
Figure 2. Structure and composition analysis of algal mats
a-b. the analysis of the microstructure of algal mats by SEM-EDS; c. the analysis of the content of C, H, N and S elements of different colors of algal mats by elemental analyzer; d. the analysis of the functional groups or chemical bonds of different colors of algal mats by FTIR; e. the analysis of the mineral phase components of algal mats by XRD
图 3 各采样点藻席的硅藻绝对丰度（Dabs）、Shannon-Weiner指数（H）、Simpson多样性指数（D）、Pielou指数（J）
Figure 3. Absolute abundance, Shannon Weiner index, Simpson's diversity index, and Pielou's species evenness index of diatoms
图 4 各样点硅藻优势种组成占比
Figure 4. Proportion of dominant species of diatom in each site in Huanglong Scenic and Historic Interest Area
图 5 采样点水环境因子的主成分分析与Spearman 相关系数
Figure 5. PCA and Spearman correlation coefficient of water environmental factors at sampling points
表 1 黄龙风景区采样点地理信息和水样物理化学参数
Table 1. Geographic information of sampling points and physicochemical parameters of water samples
编号 #1 #2 #3 #4 #5 #6 #7 #8 变异系数/% 采样点 转花泉 五彩池 争艳彩池 盆景池 潋滟湖 金沙铺地 飞瀑流辉 迎宾池 n.a. 海拔/m 3 588.00 3 574.00 3 432.00 3 328.00 3 297.00 3 239.00 3 226.00 3 220.00 n.a. 经度 103°49′28″E 103°49′58″E 103°49′51″E 103°49′44″E 103°49′37″E 103°49′29″E 103°49′36″E 103°49′25″E n.a. 纬度 32°43′28″N 32°43′30″N 32°44′23″N 32°44′47″N 32°44′50″N 32°45′1″N 32°45′2″N 32°45′4″N n.a. T/ ℃ 7.20 6.50 7.70 7.60 7.30 9.60 7.60 8.30 11.82 pH 6.78 7.50 8.10 8.44 8.57 8.40 8.49 8.50 7.87 Ec/us 1 116.00 1 075.00 627.00 523.00 507.00 515.00 510.00 501.00 39.43 pCO2/Pa 0.15 0.01 0.00 0.00 0.00 0.00 0.00 0.00 249.54 ${\rm{CO}}_3^{2-}$/mg·L−1 0.00 0.00 0.00 15.005 0.00 0.00 0.00 30.01 198.41 ${\rm{HCO}}_3^{-}$/mg·L−1 1 006.83 732.24 366.12 274.59 335.61 282.22 320.36 221.20 62.63 TN/mg·L−1 0.02 0.04 0.19 0.11 0.15 0.14 0.12 0.26 59.94 TP/mg·L−1 0.05 0.05 0.13 0.05 0.04 0.04 0.04 0.03 58.78 Ca2+ /mg·L−1 227.80 190.60 122.30 100.90 94.30 135.30 84.90 107.80 38.03 Al3+ /mg·L−1 0.06 0.06 0.04 0.03 0.03 0.01 0.03 0.03 46.48 K+/mg·L−1 0.56 0.92 0.37 0.37 0.42 0.35 0.40 0.37 41.19 Mg2+/mg·L−1 23.10 22.39 15.16 15.33 15.63 15.57 15.96 15.76 19.22 DSi/mg·L−1 2.69 2.74 2.00 1.96 1.97 1.99 1.93 1.92 16.28 Cl−/mg·L−1 0.74 1.08 0.90 0.78 0.91 0.98 0.70 0.83 14.75 ${\rm{NO}}_3^{-}$/mg·L−1 0.27 0.39 0.88 1.26 1.26 1.45 1.21 1.29 44.43 ${\rm{SO}}_4^{2-}$/mg·L−1 57.44 41.14 53.94 48.52 47.96 54.75 46.93 47.24 10.61 
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表 2 黄龙风景区藻席硅藻种类名录
Table 2. List of diatom species in the algal mats of Huanglong Scenic and Historic Interest Area
属 种 属 种 曲壳藻属（Achnanthes） 线型曲壳藻 A. orenulata 等片藻属（Diatoma） 念珠等片藻 D. moniliformis 披针形曲壳藻 A. lanceolata 冬生等片藻 D. hiemale 毛利曲壳藻 A. mauiensis 巨大等片藻 D. maximum 曲丝藻属（Achnanthidium） *极小曲丝藻 A. minutissimum 普通等片藻 D. vulgare 美壁藻属（Caloneis） 高山美壁藻 C. alpestris 中型等片藻 D. mesodon 偏肿美壁藻 C. ventricosa 纤细等片藻 D. tenue 舒曼美壁藻 C. schumanniana 双壁藻属（Diploneis） 椭圆双壁藻 D. elliptica 美丽美壁藻 C. pulchra 卵圆双壁藻长圆变种
D. ovalis var. oblongella卵形藻属（Cocconeis） 弯曲卵形藻 C. flexella 眼斑双壁藻 D. oculata 扁圆卵形藻 C. placentula 内丝藻属（Encyonema） 西里西亚内丝藻 E. silesiacum 小环藻属（Cyclotella） 可辨小环藻 C. distinguenda 簇生内丝藻 E. cespitosum 梅尼小环藻 C. meneghiniana 纤细内丝藻 E. gracile 波缘藻属（Cymatopleura） *草鞋形波缘藻 C. solea 内丝藻属（Encyonopsis） 斯托特拟内丝藻 E. stodderi 桥弯藻属（Cymbella） 新箱型桥弯藻 C. neocistula 赛萨特拟内丝藻 E. cesatii 背弯桥弯藻 C. dorsenotata 脆杆藻属（Fragilaria） 两头脆杆藻 F. biceps 近轴桥弯藻 C. proxima 狭辐节脆杆藻 F. leptostauron 桥弯藻属（Cymbella） 近淡黄桥弯藻 C. subhelvetica 异极藻属（Gomphonema） 短纹异极藻 G. abbreniatum 近平截桥弯藻 C. subtruncata *窄异极藻 G. productum *新月形桥弯藻 C. cymbiformis 直链藻属（Melosira） 变异直链藻 M. varians *箱型桥弯藻 C. cistula 颗粒直链藻极狭变种
M. granulate var.angutissima斯图施拜桥弯藻 C. stuxbergii 舟型藻属（Navicula） 隐头舟形藻 N. cryptocephala 膨胀桥弯藻 C. tumida 簇生舟形藻 N. gregaria 平滑桥弯藻 C. laevis 长篦藻属（Neidium） *二哇长篦藻 N. bisulcatum 微细桥弯藻 C. parva 优美长篦藻 N. mirabile *纤细桥弯藻 C. gracillis 菱形藻属（Nitzschia） 细长菱形藻 N. gracilis *近北极桥弯藻 C. subarctica 细端菱形藻 N. dissipata 近缘桥弯藻 C. affinis 羽纹藻属（Pinnularia） 中型羽纹藻 P. intermedia 弯肋藻属（Cymbopleura） *奥地利弯肋藻 C. austriaca 弯楔藻属（Rhoicosphenia） 弯形弯楔藻 R. curvata 库尔伯斯弯肋藻 C. kuelbsii 辐节藻属（Stauroneis） *沼泽辐节藻 S. palustris 矩圆弯肋藻 C. oblongata 十字脆杆藻属（Staurosira） 凸腹十字脆杆藻 S. venter 杂种弯肋藻 C. hybrida 针杆藻属（Synedra） 肘状针杆藻 S. ulna *窄弯肋藻 C. angustata 双头针杆藻 S. amphicephala 双头弯肋藻 C. amphicephala 尖针杆藻 S. acus *线形弯肋藻 C. linearis 平片针杆藻 S. tabulata 等片藻属（Diatoma） D. hyemalis 平板藻属（Tabellaria） 窗格平板藻 T. fenestrata *D. vulgaris 四环藻属（Tetracyclus） 湖沼四环藻 T. lacustris 注：*代表优势种。Note: * represents dominant species.
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表 3 黄龙风景区藻席硅藻优势种
Table 3. Dominant species of diatom in Huanglong Scenic and Historic Interest Area
属 种 优势度（Y） 曲丝藻属（Achnanthidium） 极小曲丝藻 A. minutissimum 0.039 波缘藻属（Cymatopleura） 草鞋形波缘藻 C. solea 0.045 桥弯藻属（Cymbella） 箱型桥弯藻 C. cistula 0.027 新月形桥弯藻 C. cymbiformis 0.030 近北极桥弯藻 C. subarctica 0.032 纤细桥弯藻 C. gracillis 0.043 弯肋藻属（Cymbopleura） 线形弯肋藻 C. linearis 0.023 窄弯肋藻 C. angustata 0.025 奥地利弯肋藻 C. austriaca 0.028 等片藻属（Diatoma） D. vulgaris 0.053 异极藻属（Gomphonema） 窄异极藻 G. productum 0.035 长篦藻属（Neidium） 二哇长篦藻 N. bisulcatum 0.048 辐节藻属（Stauroneis） 沼泽辐节藻 S. palustris 0.026
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