Preliminary study on connectivity of organism communities in artificial and natural habitats
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摘要: 为了深入了解近岸海域不同生境间的群落格局及其相互联系，有必要探究影响人工生境和自然生境物种分布与生态连通性的潜在因素。本研究对江苏海州湾四种典型生境——人工鱼礁区（ARA）、紫菜养殖区（NCA）、牡蛎礁区（OCA）、天然海区（NA）的鱼类和无脊椎动物群落开展综合调查。结果显示，四种生境的物种丰度存在显著差异（P<0.05），其中有一些重要物种能够同时存在于多种生境，也有些特有物种仅存于一种生境内。ARA中的口虾蛄（Oratosquilla oratoria）生物量远高于其它生境，矛尾虾虎鱼体长在ARA、NCA和OCA均显著大于NA（P<0.05），这一差异与饵料生物分布密切相关。不同生活习性的鱼类体长分布差异表明，物种洄游行为在生物分布和生境连通中发挥重要作用。洄游性鱼类小黄鱼（Larimichthys polyactis）和青鳞小沙丁鱼（Sardinella zunasi）在人工生境内的体长显著大于自然生境（P<0.05）。定栖性鱼类如矛尾虾虎鱼（Chaeturichthys stigmatias）体长在OCA显著大于ARA和NCA（P<0.05），而焦氏舌鳎（Cynoglossus joyneri）体长在NCA显著大于ARA和OCA（P<0.05）。本研究表明：人工生境通过提升生境复杂性，形成有利于海洋生物资源恢复和个体发育的环境条件。饵料生物分布和物种洄游特性与不同生境间的连通性具有潜在的联系。Abstract: To better understand the community patterns and their interconnections between different habitats in coastal areas, it is essential to explore the potential factors influencing species distribution and ecological connectivity between artificial and natural habitats. This study conducted a comprehensive survey of fish and invertebrate communities across four typical habitats in Haizhou Bay, Jiangsu Province: artificial reef area (ARA), nori culture area (NCA), oyster culture area (OCA), and natural area (NA). The results showed significant differences in species abundance among the four habitats (P<0.05), with some important species occurring across multiple habitats, and some endemic species restricted to a single habitat. The biomass of Oratosquilla oratoria in ARA was significantly higher compared to other habitats, and the body length of Chaeturichthys stigmatias in ARA, NCA, and OCA was significantly greater than in NA (P<0.05), which was strongly associated with the distribution of prey organisms. Differences in body length distributions of fish with different life habits indicate that species migration behavior plays an important role in species distribution and habitat connectivity. Migratory fish species, such as Larimichthys polyactis and Sardinella zunasi, exhibited significantly greater body lengths in artificial habitats compared to natural habitats (P<0.05). Resident fish species, such as Chaeturichthys stigmatias, exhibited significantly larger body lengths in OCA compared to ARA and NCA (P<0.05), while Cynoglossus joyneri exhibited a significantly greater body length in NCA than in ARA and OCA (P<0.05). This study demonstrates that artificial habitats, by enhancing habitat complexity, provide favorable environmental conditions for the restoration of marine biological resources and the development of individuals. The distribution of prey organisms and species migration characteristics are likely associated with connectivity between different habitats.
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Key words:
- Haizhou Bay /
- artificial reef area /
- oyster culture area /
- nori culture area /
- natural area /
- organism community
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图 1 江苏省海州湾研究区域和采样点
ARA、NCA、OCA和NA分别代表人工鱼礁区、紫菜养殖区、牡蛎礁区和天然海区
Fig. 1 Study area and sampling point of Haizhou Bay, Jiangsu Province
ARA, NCA, OCA and NA represent artificial reef area, nori culture area, oyster culture area and natural area, respectively
图 2 四种生境中鱼类、虾类、蟹类、头足类、贝类和棘皮类的种类数量（a）与个体数量（b）
误差条表示每组中单个数字的标准误差。下同
Fig. 2 Number of species (a) and individuals (b) of fish, shrimp, crab, cephalopod, shellfish and echinoderm in the four habitats
Error bars indicate the standard error of individual numbers in each group. Same below
图 3 四种生境中不同科的种类数量（a）与个体数量（b）
Fig. 3 Number of species (a) and individuals (b) of different families in four habitats
图 4 海州湾不同生境类型生物群落组成的非度量多维尺度（NMDS）的可视化结果。
胁强系数（0.1956<0.2）表明了排序模型的合理性。不同颜色的点代表不同的生境类型，每个点代表一个采样点
Fig. 4 Visualisation results of the non-metric multidimensional scaling (NMDS) of biotope composition of different habitat types in Hai Zhou Bay
The coefficient of coercion (0.1956<0.2) indicates the reasonableness of the ordination model. Different coloured dots represent different habitat types and each dot represents a sampling point
图 5 物种在四种生境类型中的聚类热图
显示了66个（左侧聚类）物种及其生境关联，从红色（强相关）到白色（弱相关）的颜色梯度表示物种与生境关联的强度
Fig. 5 Heat map of species clustering in four habitat types
The figure shows 66 (clustered on the left) species and their habitat associations, with a colour gradient from red (strongly associated) to white (weakly associated) indicating the strength of the species' association with the habitat
图 6 海州湾多生境关联的11种丰度较大物种的不同体长分布
方框的上限和下限分别表示第一和第三分位数，框的内线表示数据分布的平均值。（A. 赤鼻棱鳀, B. 青鳞小沙丁鱼, C. 小黄鱼, D. 皮氏叫姑鱼, E. 矛尾虾虎鱼, F. 红狼牙虾虎鱼, G. 日本蟳, H. 口虾蛄, I. 焦氏舌鳎, J. 鲜明鼓虾, K. 火枪乌贼）。注释：“*”、“**”表示存在显著差异，“***”表示存在极显著差异
Fig. 6 Distribution of different body lengths of 11 species with high abundance in multi-habitat associations in Hai Zhou Bay
The upper and lower limits of the boxes represent the first and third quartiles, respectively, and the inner line of the box represents the mean of the data distribution. (A. Thryssa kammalensis, B. Sardinella zunasi, C. Larimichthys polyactis, D. Johnius belangerii, E. Chaeturichthys stigmatias, F. Odontamblyopus rubicundus, G. Charybdis japonica, H. Oratosquilla oratoria, I. Cynoglossus joyneri, J. Alpheus distinguendus, and K. Loligo beka). Note: "*" and "**" indicate significant differences, and "***" indicates highly significant differences
表 1 生境类型及组成特点
Tab. 1 Habitat types and composition characteristics
生境类型 组成特点 紫菜养殖区 主要使用浮筏装置养殖紫菜 人工鱼礁区 由一系列人工鱼礁(如混凝土立方礁、十字形礁、塔形礁)组成的人工鱼礁区 天然海区 未受明显人为干扰的自然海域 牡蛎礁区 由活体牡蛎、死亡牡蛎的壳及其他礁区生物共同堆积组成的聚集体组成的生态修复区 
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表 2 四种生境中鱼类优势种和重要种的相对重要性指数
Tab. 2 Relative importance indices of dominant and important species of fish in four habitats
物种种类 拉丁名 IRI值 矛尾虾虎鱼 Chaeturichthys stigmatias 3191.24 赤鼻棱鳀 Thryssa kammalensis 2270.09 小黄鱼 Larimichthys polyactis 1216.64 斑鰶 Konosirus punctatus 889.74 焦氏舌鳎 Cynoglossus joyneri 840.05 青鳞小沙丁鱼 Sardinella zunasi 606.95 皮氏叫姑鱼 Johnius belangerii 552.97 黄鲫 Setipinna tenuifilis 487.22 红狼牙虾虎鱼 Odontamblyopus rubicundus 455.66 香斜棘䲗 Repomucenus olidus 307.18 白姑鱼 Pennahia argentata 284.60 条鲾 Equulites rivulatus 260.91 银鲳 Pampus argenteus 214.79 半滑舌鳎 Cynoglossus semilaevis 213.04 鲬 Platycephalus indicus 169.42 中华栉孔虾虎鱼 Ctenotrypauchen chinensis 165.96
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表 3 四种生境的鱼类生物多样性
Tab. 3 Fish biodiversity in four habitats
站点 H'值 J值 D值 ARA-1 1.99 0.64 3.19 ARA-2 1.52 0.56 2.11 ARA-3 1.97 0.63 3.23 ARA-4 2.64 0.80 3.88 NCA-1 2.36 0.83 3.29 NCA-2 1.56 0.87 0.95 NCA-3 1.41 0.73 1.12 NCA-4 1.06 0.44 1.86 OCA-1 1.63 0.71 1.88 OCA-2 1.80 0.68 2.15 OCA-3 1.87 0.71 2.35 OCA-4 1.87 0.68 2.82 NA-1 0.69 0.29 1.51 NA-2 0.91 0.40 1.53 NA-3 1.63 0.71 1.93 NA-4 2.49 0.92 2.89
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表 4 PERMANOVA 组间比较结果
Tab. 4 PERMANOVA intergroup comparison results
Df Sums of
squaresMean
squaresF.Model Variation
(R2)Pr
(>F)ARA/NCA 1 0.462 0.462 1.569 20.73% 0.095 ARA/OCA 1 0.366 0.366 1.148 14.08% 0.285 ARA/NA 1 0.523 0.523 1.495 15.74% 0.016 NCA/OCA 1 0.542 0.542 1.578 23.99% 0.108 NCA/NA 1 0.597 0.597 1.570 20.74% 0.055 OCA/NA 1 0.331 0.331 0.841 10.73% 0.718
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