Spatio-temporal niches and interspecific association of dominant fish species in the offshore waters of southern Zhejiang
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摘要: 基于2016年8月（夏季）、11月（秋季）、2017年3月（冬季）、5月（春季）浙江南部近海的鱼类调查数据，利用生态位宽度、生态位重叠、聚类分析、方差比率法、卡方检验、联结系数以及种对共同出现百分率对主要鱼类种间关系进行研究。结果表明：（1）调查共采集鱼类169种，主要鱼类共有26种；（2）在空间维度上，日本鲭（Scomber japonicus）生态位宽度最大（2.83），青鳞小沙丁（Sardinella zunasi）和芝芜棱鳀（Thryssa chefuensis）生态位重叠值最大（0.97）；在时间维度上，龙头鱼（Harpadon nehereus）生态位宽度最大（1.34），6组种对时间生态位重叠值等于1.00；在时空维度上，龙头鱼生态位宽度最大（3.50），青鳞小沙丁与芝芜棱鳀时空生态位重叠值最大（0.97）；（3）方差比率法分析表明，方差比率偏离显著，主要鱼类总体正联结显著，其中170组种对间联结性达到显著水平（
$ \chi^2$ ≥3.841），联结系数和种对共同出现频率结果表明种间联结性趋于正联结。Abstract: Based on the fish survey data of the offshore waters of southern Zhejiang in August 2016 (summer), November 2016 (Autumn), March 2017 (winter) and May 2017 (spring), the relationship between dominant fish species is determined by niche breadth, niche overlap, cluster analysis, variance ratio (VR) method, chi-square test, association coefficient (AC) and species pair common percentage (PC). The results show that: (1) 169 species of fish are collected, including 26 major species; (2) in the spatial dimension, the niche breadth of Scomber japonicus is the largest (2.83); the niche overlap between Sardinella zunasi and Thryssa chefuensis is the largest (0.97); in the temporal dimension, the niche breadth of Harpadon nehereus is the largest (1.34); the niche overlap of six groups is equal to 1.00; in the spatial and temporal dimension, the niche breadth of Harpadon nehereus is the largest (3.08); the niche overlap between Sardinella zunasi and Thryssa chefuensis is the largest (0.97); (3) the variance ratio analysis showed that the VR deviation is significant, and the overall positive association of the major fish is significant, among which 170 groups of species pair association reached a significant level (χ2≥3.841). The results of the AC and the PC showed that the interspecific connection tended to be positive. -
图 2 浙江南部近海主要鱼类空间生态位宽度相似性聚类分析
Fig. 2 Cluster analysis of spatial niche breadth similarity of dominant fish species in the offshore waters of southern Zhengjiang
图 3 浙江南部近海主要鱼类时间生态位宽度相似性聚类分析
Fig. 3 Cluster analysis of temporal niche breadth similarity of dominant fish species in the offshore waters of southern Zhengjiang
图 4 浙江南部近海主要鱼类时空生态位宽度相似性聚类分析
Fig. 4 Cluster analysis of spatial-temporal niche breadth similarity of dominant fish species in the offshore waters of southern Zhengjiang
图 5 浙江南部近海主要鱼类种间
$ \chi^2$ 检验半矩阵Fig. 5 Semi-matrix of the inters
$ \chi^2$ test among the dominant fish species in the offshore waters of southern Zhengjiang
图 6 浙江南部近海主要鱼类种间联结系数（AC）半矩阵
Fig. 6 Semi-matrix of inters pecific association coefficient (AC) among the dominant fish species in the offshore waters of southern Zhengjiang
图 7 浙江南部近海主要鱼类种间共同出现百分率(PC)半矩阵
Fig. 7 Semi-matrix of inters percentage of co-occurrence (PC) among the dominant fish species in the offshore waters of southern Zhengjiang
表 1 浙江南部近海鱼类重要种种类组成
Tab. 1 Major composition of dominant fish species in the offshore waters of southern Zhengjiang
种号 种类 重要性指数IRI 春季 夏季 秋季 冬季 1 刺鲳Psenopsis anomala 486.70 1 966.84 255.20 − 2 带鱼Trichiurus lepturus 3 686.91 9 790.40 1 792.75 421.48 3 黑姑鱼Atrobucca nibe 516.24 1 684.04 − − 4 镰鲳Pampus echinogaster 244.47 861.85 199.22 − 5 龙头鱼Harpadon nehereus 2 307.57 928.74 8 000.79 2 774.79 6 小黄鱼Larimichthys polyactis 200.64 1 026.15 594.08 748.10 7 二长棘犁齿鲷Evynnis cardinalis − 138.09 − − 8 黄鲫Setipinna tenuifilis 5 970.94 458.82 289.89 156.51 9 竹荚鱼Trachurus japonicus − 324.02 − − 10 银姑鱼Pennahia argentata − 362.68 − − 11 六丝钝尾虾虎鱼Amblychaeturichthys hexanema − − 1 410.23 − 12 细条天竺鲷Apogonichthys lineatus 156.61 − 950.65 106.24 13 鳞鳍叫姑鱼Johnius distinctus 119.95 − 160.20 − 14 鮸Miichthys miiuy − − 152.14 − 15 皮氏叫姑鱼Johnius belangerii − − 200.81 − 16 前肛鳗Dysomma anguillare − − 178.97 − 17 沙带鱼Lepturacanthus savala − − 125.36 − 18 芝芜棱鳀Thryssa chefuensis − − − 10 284.14 19 七星底灯鱼Benthosema pterotum − − − 491.30 20 拟尖鳍犀鳕Bregmaceros pseudolanceolatus 114.11 − 288.42 186.82 21 黄鮟鱇Lophius litulon − − − 177.47 22 青鳞小沙丁Sardinella zunasi − − − 198.70 23 日本鲭Scomber japonicus 1 724.15 − − − 24 蓝圆鲹Decapterus maruadsi 365.30 − − − 25 大头银姑鱼Pennahia macrocephalus − − 162.94 − 26 日本鳀Engraulis japonicus 400.55 − − − 注：表中“−”代表物种IRI小于100，并非该季节重要种。 
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表 2 浙江南部近海主要鱼类的空间、时间及时空生态位宽度
Tab. 2 Spatial, temporal and spatio-temporal niche breadth of dominant fish species in the offshore waters of southern Zhengjiang
种类 空间生态位宽度 时间生态位宽度 时空生态位宽度 刺鲳Psenopsis anomala 2.47 0.35 0.87 带鱼Trichiurus lepturus 2.50 0.40 1.01 黑姑鱼Atrobucca nibe 2.56 0.21 0.54 镰鲳Pampus echinogaster 2.59 0.57 1.47 龙头鱼Harpadon nehereus 2.61 1.34 3.50 小黄鱼Larimichthys polyactis 2.53 1.00 2.53 二长棘犁齿鲷Evynnis cardinalis 1.79 0.22 0.39 黄鲫Setipinna tenuifilis 2.48 0.91 2.26 竹荚鱼Trachurus japonicus 2.12 0.39 0.82 银姑鱼Pennahia argentata 2.14 0.55 1.18 六丝钝尾虾虎鱼Amblychaeturichthys hexanema 2.62 0.71 1.87 细条天竺鲷Apogonichthys lineatus 2.04 1.15 2.35 鳞鳍叫姑鱼Johnius distinctus 2.43 1.01 2.44 鮸Miichthys miiuy 2.45 0.63 1.55 皮氏叫姑鱼Johnius belangerii 2.59 1.14 2.95 前肛鳗Dysomma anguillare 2.44 1.21 2.95 沙带鱼Lepturacanthus savala 2.45 0.78 1.92 芝芜棱鳀Thryssa chefuensis 0.84 0.04 0.03 七星底灯鱼Benthosema pterotum 1.87 0.91 1.70 拟尖鳍犀鳕Bregmaceros pseudolanceolatus 1.90 0.95 1.81 黄鮟鱇Lophius litulon 2.31 0.15 0.34 青鳞小沙丁Sardinella zunasi 0.54 0.10 0.05 日本鲭Scomber japonicus 2.83 0.29 0.82 蓝圆鲹Decapterus maruadsi 2.55 0.93 2.38 大头银姑鱼Pennahia macrocephalus 2.21 0.79 1.75 日本鳀Engraulis japonicus 1.30 0.46 0.60
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表 3 浙江南部近海主要鱼类空间生态位（对角线以下）和时间生态位（对角线以上）重叠值
Tab. 3 Temporal niche overlaps (above the diagonal) and spatial niche overlaps (under the diagonal) between dominant fish species in the offshore waters of southern Zhejiang
种号 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 1 1.00 0.999 0.999 0.60 0.89 0.999 0.48 0.998 0.999 0.13 0.09 0.30 0.04 0.31 0.56 0.06 0.001 0.03 0.04 0.01 0.01 0.13 0.56 0.98 0.05 2 0.81 0.999 0.999 0.61 0.90 0.999 0.48 0.998 0.999 0.11 0.09 0.29 0.04 0.30 0.55 0.05 0.02 0.04 0.06 0.03 0.03 0.13 0.56 0.98 0.05 3 0.63 0.66 0.997 0.58 0.89 0.999 0.46 0.997 0.997 0.10 0.06 0.27 0.02 0.28 0.54 0.03 0.001 0.01 0.03 0.004 0.01 0.11 0.53 0.97 0.02 4 0.71 0.61 0.44 0.63 0.90 0.997 0.52 0.998 1.00 0.15 0.13 0.34 0.07 0.34 0.59 0.09 0.02 0.06 0.08 0.03 0.03 0.17 0.59 0.99 0.09 5 0.31 0.37 0.29 0.32 0.85 0.58 0.54 0.60 0.63 0.61 0.83 0.61 0.71 0.77 0.85 0.61 0.61 0.77 0.77 0.62 0.62 0.30 0.58 0.69 0.34 6 0.82 0.73 0.78 0.73 0.29 0.89 0.47 0.89 0.91 0.24 0.42 0.32 0.31 0.44 0.64 0.19 0.44 0.46 0.48 0.45 0.45 0.12 0.52 0.89 0.11 7 0.42 0.48 0.46 0.38 0.30 0.39 0.49 0.999 0.998 0.09 0.06 0.28 0.002 0.27 0.53 0.02 0.00 0.01 0.03 0.004 0.01 0.14 0.56 0.98 0.05 8 0.33 0.56 0.37 0.29 0.45 0.17 0.45 0.53 0.51 0.17 0.42 0.84 0.07 0.38 0.49 0.32 0.06 0.29 0.29 0.09 0.07 0.93 0.99 0.60 0.90 9 0.63 0.43 0.55 0.32 0.25 0.63 0.42 0.13 0.999 0.10 0.09 0.32 0.01 0.29 0.55 0.04 0.002 0.03 0.05 0.01 0.01 0.19 0.60 0.98 0.11 10 0.48 0.59 0.50 0.18 0.34 0.23 0.33 0.66 0.16 0.14 0.12 0.33 0.06 0.33 0.57 0.08 0.03 0.06 0.08 0.04 0.04 0.16 0.58 0.99 0.09 11 0.57 0.45 0.53 0.50 0.48 0.52 0.17 0.29 0.29 0.41 0.77 0.65 0.93 0.97 0.88 0.97 0.08 0.38 0.37 0.08 0.09 0.10 0.28 0.27 0.12 12 0.54 0.61 0.71 0.59 0.14 0.79 0.10 0.04 0.32 0.10 0.58 0.70 0.90 0.85 0.77 0.84 0.62 0.86 0.86 0.63 0.63 0.37 0.44 0.23 0.46 13 0.41 0.19 0.41 0.23 0.43 0.22 0.22 0.35 0.15 0.56 0.63 0.11 0.51 0.77 0.77 0.78 0.01 0.36 0.35 0.02 0.01 0.80 0.88 0.48 0.80 14 0.22 0.23 0.27 0.19 0.62 0.29 0.07 0.24 0.20 0.29 0.42 0.17 0.56 0.91 0.80 0.90 0.44 0.67 0.66 0.44 0.45 0.001 0.15 0.16 0.08 15 0.60 0.43 0.44 0.66 0.43 0.57 0.16 0.19 0.30 0.43 0.80 0.48 0.61 0.52 0.96 0.96 0.18 0.49 0.48 0.19 0.19 0.25 0.48 0.46 0.28 16 0.53 0.47 0.62 0.38 0.58 0.67 0.20 0.14 0.66 0.22 0.43 0.52 0.40 0.55 0.47 0.86 0.16 0.44 0.43 0.16 0.17 0.28 0.60 0.68 0.28 17 0.19 0.32 0.24 0.21 0.76 0.17 0.13 0.56 0.11 0.47 0.52 0.04 0.48 0.79 0.48 0.41 0.11 0.45 0.43 0.12 0.12 0.30 0.41 0.22 0.34 18 0.03 0.04 0.04 0.12 0.54 0.13 0.01 0.05 0.01 0.06 0.17 0.02 0.45 0.67 0.33 0.59 0.55 0.93 0.93 1.00 1.00 0.003 0.001 0.004 0.13 19 0.27 0.40 0.67 0.14 0.28 0.43 0.31 0.31 0.17 0.51 0.20 0.27 0.28 0.16 0.22 0.45 0.24 0.09 1.00 0.93 0.93 0.24 0.25 0.10 0.36 20 0.17 0.26 0.19 0.19 0.12 0.19 0.03 0.08 0.30 0.14 0.20 0.19 0.13 0.25 0.30 0.38 0.20 0.04 0.37 0.94 0.94 0.24 0.25 0.11 0.36 21 0.33 0.35 0.21 0.20 0.37 0.28 0.07 0.23 0.13 0.25 0.29 0.16 0.35 0.58 0.41 0.28 0.40 0.29 0.14 0.21 1.00 0.03 0.02 0.01 0.15 22 0.02 0.01 0.02 0.10 0.49 0.13 0.001 0.04 0.001 0.002 0.07 0.01 0.41 0.63 0.20 0.59 0.46 0.97 0.09 0.003 0.28 0.004 0.01 0.01 0.13 23 0.64 0.64 0.74 0.57 0.54 0.71 0.29 0.39 0.27 0.39 0.65 0.66 0.54 0.47 0.55 0.61 0.41 0.31 0.50 0.15 0.39 0.31 0.88 0.27 0.99 24 0.47 0.52 0.44 0.53 0.36 0.30 0.79 0.60 0.34 0.43 0.39 0.13 0.45 0.19 0.36 0.19 0.31 0.05 0.19 0.18 0.28 0.02 0.43 0.68 0.84 25 0.29 0.52 0.34 0.20 0.41 0.12 0.18 0.77 0.08 0.78 0.35 0.04 0.37 0.47 0.42 0.16 0.71 0.14 0.34 0.23 0.40 0.04 0.30 0.41 0.20 26 0.08 0.09 0.08 0.17 0.07 0.10 0.04 0.11 0.16 0.01 0.10 0.09 0.01 0.05 0.09 0.12 0.01 0.01 0.05 0.10 0.07 0.02 0.35 0.29 0.02 注：该表种号同表1；表中黑体数字为生态位重叠值大于0.60的两种生物，表示生态位重叠程度较高。
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表 4 浙江南部近海主要鱼类时空生态位重叠值
Tab. 4 Spatio-temporal niche overlap between dominant fish species in the offshore waters of southern Zhejiang
种号 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 2 0.81 3 0.63 0.66 4 0.71 0.61 0.44 5 0.18 0.23 0.17 0.20 6 0.73 0.65 0.70 0.66 0.25 7 0.42 0.48 0.46 0.38 0.17 0.35 8 0.16 0.27 0.17 0.15 0.24 0.08 0.22 9 0.62 0.43 0.55 0.32 0.15 0.56 0.42 0.07 10 0.48 0.59 0.50 0.18 0.21 0.21 0.33 0.34 0.16 11 0.07 0.05 0.05 0.08 0.29 0.13 0.01 0.05 0.03 0.06 12 0.05 0.06 0.04 0.08 0.12 0.33 0.01 0.02 0.03 0.01 0.45 13 0.12 0.05 0.11 0.08 0.26 0.07 0.06 0.29 0.05 0.18 0.41 0.07 14 0.01 0.01 0.01 0.01 0.44 0.09 <0.001 0.02 0.002 0.02 0.38 0.16 0.29 15 0.19 0.13 0.13 0.22 0.33 0.25 0.04 0.07 0.09 0.14 0.78 0.41 0.47 0.47 16 0.30 0.26 0.33 0.22 0.49 0.43 0.11 0.07 0.36 0.12 0.37 0.40 0.31 0.43 0.45 17 0.01 0.01 0.01 0.02 0.46 0.03 0.003 0.18 0.005 0.04 0.51 0.04 0.38 0.71 0.46 0.35 18 <0.001 0.001 <0.001 0.003 0.33 0.06 0.00 <0.001 <0.001 0.002 0.01 0.01 0.002 0.30 0.06 0.09 0.06 19 0.01 0.02 0.01 0.01 0.21 0.20 0.004 0.09 0.01 0.03 0.07 0.24 0.10 0.11 0.11 0.20 0.11 0.08 20 0.01 0.01 0.01 0.01 0.09 0.09 0.001 0.02 0.01 0.01 0.07 0.17 0.05 0.16 0.14 0.17 0.09 0.03 0.37 21 0.002 0.01 0.001 0.01 0.22 0.12 <0.001 0.02 0.001 0.01 0.02 0.10 0.01 0.26 0.08 0.04 0.05 0.29 0.13 0.20 22 <0.001 <0.001 <0.001 0.003 0.30 0.06 <0.001 0.003 <0.001 <0.001 0.01 0.01 0.005 0.28 0.04 0.10 0.05 0.97 0.08 0.003 0.28 23 0.08 0.08 0.08 0.10 0.16 0.09 0.04 0.36 0.05 0.06 0.06 0.25 0.44 <0.001 0.14 0.17 0.12 0.001 0.12 0.04 0.01 0.001 24 0.26 0.29 0.23 0.32 0.21 0.16 0.44 0.59 0.20 0.25 0.11 0.06 0.39 0.03 0.17 0.12 0.13 <0.001 0.05 0.05 0.01 <0.001 0.38 25 0.29 0.51 0.33 0.20 0.28 0.10 0.17 0.46 0.08 0.76 0.09 0.01 0.18 0.08 0.19 0.11 0.16 0.001 0.03 0.03 0.004 <0.001 0.08 0.28 26 0.004 0.005 0.002 0.02 0.02 0.01 0.002 0.10 0.02 0.001 0.01 0.04 0.01 0.004 0.02 0.03 0.004 0.001 0.02 0.04 0.01 0.002 0.35 0.25 0.004 注：该表种号同表1；表中黑体数字为生态位重叠值大于0.60的两种生物，表示生态位重叠程度较高。
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表 5 浙江南部近海主要鱼类总体联结性
Tab. 5 General interspecific association test of dominant fish species in the offshore waters of southern Zhengjiang
ST2 $ \delta^2_T$ VR W $\ \chi^2$[$\chi^2_{0.95}$(27), $\ \chi^2_{0.05}$(27)] 8.63 0.77 11.27 304.19 (16.15, 40.11)
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