Analysis of invertebrate diversity and co-occurrence network based on environmental DNA metabarcoding in autumn typical tidal creek units of the Huanghe River Delta
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摘要: 潮沟系统作为滨海湿地中活跃的地貌单元，不同级别潮沟水文环境变化显著，进而导致生物群落的空间分布出现差异。本研究选取黄河三角洲典型潮沟单元，利用环境DNA宏条形码（environmental DNA metabarcoding，eDNA）技术检测不同级别潮沟无脊椎动物多样性，采用生物共现网络分析和冗余分析（RDA）分别揭示典型潮沟无脊椎动物关键种与驱动因素。结果表明：该潮沟单元共检测到无脊椎动物127个分类操作单元（Operational Taxonomic Units，OTUs），隶属于9门24纲53目103科87属90种。无脊椎动物门水平和属水平分别以节肢动物门（43.9%）和围沙蚕属（Perinereis ）（25.2%）为优势类群。群落综合多样性指标（Comprehensive diversity index，CD）分析显示，三级潮沟综合多样性最高，一级潮沟无脊椎动物综合多样性最低。生物共现网络分析显示，线围沙蚕（Perinereis linea）和双枝薮枝螅（Obelia dichotoma）为关键种，对维持该潮沟无脊椎动物群落结构稳定起关键作用。RDA显示，水体的硅酸盐含量、温度和沉积物的粉砂、黏土占比是影响该潮沟无脊椎动物群落特征的主要环境因子。相关性网络分析显示，关键种受硅酸盐含量、黏土和水体氮元素含量的显著影响（P < 0.05）。研究结果有助于了解黄河三角洲典型潮沟无脊椎动物群落结构，揭示典型潮沟无脊椎动物关键种，并为无脊椎动物多样性监测与保护提供数据支持与理论参考。Abstract: The tidal creek system is an active geomorphic unit in coastal wetlands, and the water environment of different level tidal creeks changes significantly, leading to spatial distribution differences of biological communities. This study selected a typical tidal creek unit in the Huanghe River Delta and used environmental DNA metabarcoding (eDNA) technique to detect the diversity of invertebrates. The biological co-occurrence network analysis was used to reveal the key species and driving factors of invertebrates in the typical tidal creek. The results showed that a total of 127 operational taxonomic units (OTUs) of invertebrates were detected in the tidal creek unit, belonging to 9 phyla, 24 classes, 53 orders, 103 families, 87 genera, and 90 species; among them, the class level was dominated by the Arthropoda (43.9%), and the genus level was dominated by the Perinereis (25.2%). The comprehensive diversity index (CD) analysis showed that the comprehensive diversity of invertebrates in the third-level tidal creek was the highest, and the comprehensive diversity of invertebrates in the first-level tidal creek was the lowest. The biological co-occurrence network analysis showed that the Perinereis linea and the Obelia dichotoma were the keystone species, which played a key role in maintaining the stability of the invertebrate community structure in the tidal creek. The RDA showed that the silicate content of the water body, temperature, and the proportion of fine sand and clay in the sediment were the main environmental factors affecting the invertebrate community characteristics in the tidal creek. Correlation network analysis showed that the keystone species were significantly affected by silicate content, clay, and nitrogen content in water (P < 0.05). The research results are helpful for understanding the community structure of typical tidal ditch invertebrates, revealing the keystone species of typical tidal ditch invertebrates, and providing data support and theoretical reference for the monitoring and protection of invertebrate diversity.
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图 2 黄河三角洲典型潮沟门（a）和属（b）水平无脊椎动物相对丰度组成
Fig. 2 Relative abundance of invertebrates on phylum (a) and genus (b) level in Huanghe River Delta typical tidal creek
图 3 不同采样点无脊椎动物OTUs多集韦恩图（a）与HCA（b）
Fig. 3 Multiset Venn diagram of invertebrate OTUs among different sampling sites (a) and HCA (b)
图 4 无脊椎动物科水平（a）和种水平（b）的生物共现网络拓扑图
图中节点表示无脊椎动物；不同节点颜色表示不同模块；节点大小表示物种相对丰度；边颜色表示正负相关性，红色表示正相关，绿色表示负相关；边大小表示相关性系数绝对值大小
Fig. 4 Topology of biological co-occurrence networks at family (a) and species levels (b) of invertebrates
The nodes in the graph represent invertebrates; different node colors represent different modules; node size represents the relative abundance of species. Edge colors represent positive and negative correlations, with red indicating positive correlation and green indicating negative correlation; edge size represents the absolute value of the correlation coefficient
图 5 黄河三角洲潮沟无脊椎动物门水平与环境因子的冗余分析
Fig. 5 Redundancy analysis between invertebrates on phylum level and environmental factors at Yellow River delta tidal creek
图 6 科水平（a）和种水平（b）关键类群与环境因子相关性分析
图中蓝色节点表示无脊椎动物，节点大小表示无脊椎动物相对丰度大小；红色节点表示环境因子；边颜色表示正负相关性，红色表示正相关，黑色表示负相关；边大小表示相关性系数绝对值大小
Fig. 6 Correlation analysis of keystone groups and environmental factors at family level (a) and species level (b)
The blue nodes in the figure represent invertebrates, and the size of the nodes indicates the relative abundance of invertebrates. The red nodes represent environmental factors. The color of the edges indicates the sign of the correlation, with red indicating positive correlation and black indicating negative correlation; the size of the edges indicates the absolute value of the correlation coefficient
A1 基于环境DNA宏条形码技术的典型潮沟无脊椎动物物种名录
A1 Typical Invertebrate species list of intertidal scours based on environmental DNA metabarcoding
门 属 种 环节动物门Annelida 围沙蚕属Perinereis 线围沙蚕Perinereis linea 水丝蚓属Limnodrilus 水丝蚓属一种Limnodrilus sp. 丝异须虫属Heteromastus Heteromastus gusipoensis Branchinotogluma Branchinotogluma hessleri 节肢动物门Arthropoda 纺锤水蚤属Acartia 纺锤水蚤属一种Acartia sp. Carpelimus Carpelimus sp. BBCCM392-10 Chydorus Chydorus sphaericus 伪镖水蚤属Pseudodiaptomus 指状伪镖水蚤Pseudodiaptomus inopinus 纹藤壶属Amphibalanus 象牙纹藤壶Amphibalanus eburneus 小绿叶蝉属Empoasca 小绿叶蝉属一种Empoasca sp. BOLD-2016 Eupodes Eupodes sp. BIOUG25165-F12 单蛛属Haplodrassus 单蛛属一种Haplodrassus sp. 人形鱼虱属Lernanthropus Lernanthropus corniger 湖水虱属Ligidium 湖水虱属一种Ligidium sp. 蜞属Gaetice 平背蜞Gaetice depressus 侧沟茧蜂属Microplitis 侧沟茧蜂属一种Microplitis sp. 矮胖猛水蚤属Nannopus Nannopus parvus Oxidus Oxidus sp. 拳蟹属Pyrhila 豆形拳蟹Pyrhila pisum Troglarmadillo Troglarmadillo sp. 若甲螨属Zygoribatula 若甲螨属一种Zygoribatula sp. Pleuromamma Pleuromamma abdominalis 球蛛属Coleosoma 球蛛属一种Coleosoma sp. Haloniscus Haloniscus sp. MG1641 Herbita Herbita sp. 1YB 小家蚁属Monomorium 小家蚁属一种Monomorium sp. 刺胞动物门Cnidaria Aurelia Aurelia sp. Abietinaria Abietinaria variabilis 美螅水母属Clytia 美螅水母属一种Clytia sp. WA01 哈钟螅属Hartlaubella Hartlaubella gelatinosa 水螅属Hydra 水螅属一种Hydra sp. Malagazzia 带拟杯水母Malagazzia taeniogonia Gonionemus 钩手水母属一种Gonionemus sp. 薮枝螅水母属Obelia Obelia dichotoma 薮枝螅水母属一种Obelia sp. MZUSP 3356 Obelia longissima Obelia geniculata 僧帽水母属Physalia 僧帽水母属一种Physalia sp. DRP-2009 海榧属Plumularia 海榧属一种Plumularia sp. Praya Praya dubia Thecocodium Thecocodium quadratum Zanclea Zanclea implexa 离翼水母属Apolemia 离翼水母属一种Apolemia sp. 3 CWD-2005 Caryophyllia Caryophyllia sp. 逼螅水母属Pennaria Pennaria disticha 棘皮动物门Echinodermata 长棘海星属Acanthaster Acanthaster planci 海盘车属Asterias 多棘海盘车Asterias amurensis 软体动物门Mollusca 彩虹蛤属Iridona 彩虹蛤Iridona iridescens 德文蛤属Devonia 内壳德文蛤Devonia semperi Bosellia Bosellia mimetica Chosenelix Chosenelix sp. Cornu Cornu aspersum 冠蚌属Cristaria Cristaria plicata 砂螺属Gastrocopta 砂螺属一种Gastrocopta sp. Glyptophysa Glyptophysa sp. 6 PGA-2018 Granulomelon Granulomelon sp. A FC-2015 Magallana 长牡蛎Crassostrea gigas Indoplanorbis Indoplanorbis exustus Lacuna Lacuna variegata 旋螺属Gyraulus 旋螺属一种Gyraulus sp. 小贻贝属Mytella Mytella charruana 蜑螺属Nerita Nerita magdalenae Phestilla Phestilla sp. Radix 折叠萝卜螺Radix plicatula Runcina Runcina sp. c AKA-2020 Setobaudinia Setobaudinia kessneri Trapania Trapania orteai 海蛞蝓属Tritonia 海蛞蝓属一种Tritonia sp. Clathurella Daphnella sp. NP-2008 Digidentis Digidentis cf. arbutus LMT-2007 Fissurella Fissurella bravensis Kermia Kermia melanoxytum 叶海牛属Phyllidiella Phyllidiella albonigra 纽形动物门Nemertea 四眼属Tetrastemma 四眼属一种Tetrastemma sp. 线虫动物门Nematoda 阔口线虫属Eurystomina 阔口线虫属一种Eurystomina sp. 1AMA Obainia Obainia pachnephorus Adoncholaimus Adoncholaimus daikokuensis 扁形动物门Platyhelminthes Bolbophorus Bolbophorus damnificus Cratera Cratera tui 单咽虫属Haplopharynx 单咽虫属一种Haplopharynx sp. 多目涡虫属Polycelis 多目涡虫属一种Polycelis sp. Pseudoceros Pseudoceros laingensis Syndesmis Syndesmis kurakaikina 副似四钩虫属Paratetraonchoides 副似四钩虫属一种Paratetraonchoides sp. 多孔动物门Porifera Caulophacus Caulophacus arcticus 石豆海绵属Cyamon 石豆海绵属一种Cyamon sp. Scopalina Scopalina canariensis Suberites Suberites topsenti Thrinacophora Thrinacophora cervicornis Arturia 细薄白枝海绵Arturia tenuipilosa 
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表 1 DNA宏条形码测序结果
Tab. 1 environmental DNA metabarcoding sequencing results
类群 序列数 OTUs 序列百分比 节肢动物门Arthropoda 16123 50 38.4% 环节动物门Annelida 11645 6 27.8% 刺胞动物门Cnidaria 5662 22 13.5% 软体动物门Mollusca 3920 28 9.4% 线虫动物门Nematoda 2543 4 6.1% 棘皮动物门Echinodermata 716 2 1.7% 扁形动物门Platyhelminthes 704 7 1.7% 多孔动物门Porifera 426 5 1.0% 纽形动物门Nemertea 206 3 0.5% 合计 41945 127 100%
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表 2 黄河三角洲典型潮沟无脊椎动物α多样性指数
Tab. 2 Invertebrate alpha diversity index in typical tidal creek of Haunghe River Delta
样点 辛普森
指数香农
指数均匀度
指数Chao1
指数ACE
指数综合多样性
指数S3S 0.945 3.539 0.764 109.000 107.300 0.21 S2S 0.913 3.344 0.736 115.000 104.100 0.18 S2X 0.723 2.527 0.563 89.500 89.180 0.03 S1S 0.601 1.790 0.394 105.200 98.930 0.05 S1X 0.938 3.416 0.748 97.870 98.940 0.09
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表 3 无脊椎动物生物共现网络特征指标
Tab. 3 Characteristic index of invertebrate co-occurrence network
特征
指标平均度 平均加
权度网络
直径图密度 模块化
系数平均聚类
系数平均路径
长度模块数 科水平 5.92 11.38 5 0.11 0.64 0.87 1.43 3 种水平 12.84 25.69 4 0.25 0.54 0.6 2.1 8
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