Ecosystem structure in the Haizhou Bay and adjacent waters based on Ecopath model
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摘要: 基于2018年海州湾及邻近海域的渔业资源底拖网调查数据,运用Ecopath with Ecosim 6.5 (EwE)软件构建由26个功能群组成的海州湾及邻近海域生态系统Ecopath模型,对现阶段该生态系统的营养结构、营养相互关系和系统总特征等进行分析,旨在为实施基于生态系统的渔业管理提供理论依据。结果表明:海州湾及邻近海域生态系统各功能群的营养级范围为1.00~4.19,其中鱼类营养级范围较广,为3.22~4.19;浮游动物和其他软体动物受初级生产者和捕食者的双重作用,处于重要的营养位置;生态系统总体特征分析显示,该生态系统的总初级生产量与总呼吸量的比值为7.096,总初级生产量与总生物量的比值为56.866,系统的连接指数和系统杂食指数分别为0.429和0.204,说明该生态系统目前处于不成熟、不稳定的状态,容易受外界扰动的影响。本文通过对海州湾及邻近海域生态系统模型进行研究,解析了该海域营养结构和系统发育状况,将为海州湾渔业资源的可持续利用和科学管理提供理论依据。Abstract: Based on the data obtained from bottom trawl investigation in the Haizhou Bay and adjacent waters in 2018, a mass-balance model for the Haizhou Bay ecosystem was constructed by Ecopath with Ecosim software, consisting of 26 functional groups. Using Ecopath model, we evaluated the trophic structure, trophic impact relationship and ecosystem characters of the Haizhou Bay to provide a theoretical basis for the implementation of ecosystem-based fisheries management. The result showed that in this ecosystem trophic levels of functional groups varied from 1.00 to 4.19, and the range of fish trophic level was wide, ranging from 3.22 to 4.19. Phytoplankton and molluscs were in important nutritional positions in the ecosystem facing with the pressure from both the primary producers and predators. The evaluation of the ecosystem structure and function showed that the total primary production/total respiration (TPP/TR) was 7.096, total primary production /total biomass (TPP/B) was 56.866, the connectance index (CI) and system omnivory index (SOI) were 0.429 and 0.204, indicating that the Haizhou Bay ecosystem was in an immature, unstable state and was susceptible to external disturbances. Through the study of the Haizhou Bay and its adjacent waters ecosystem model, we analyzed the nutrient structure and phylogenetic status of the sea area, which would provide a theoretical basis for the sustainable utilization and scientific management of fishery resources in the Haizhou Bay.
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Key words:
- Haizhou Bay /
- Ecopath model /
- trophic structure /
- ecosystem character /
- functional group
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图 2 海州湾及邻近海域生态系统食物网
编号1~26见表1。 图中的圆圈代表功能群,其面积代表相对生物量的大小,两两间的连线代表能量传递过程
Fig. 2 Food web of the Haizhou Bay and adjacent waters
1~26 correspond numbers in Table 1. The circle in the figure represents the functional group, the area of which represents the relative biomass, and the connection between two circles represents the energy transfer process
表 1 海州湾及邻近海域生态系统Ecopath模型的功能群及主要种类组成
Tab. 1 Functional groups and main species based on Ecopath model in the Haizhou Bay and adjacent waters
编号 功能群 种类组成 1 浮游动物食性鱼类(Zooplanktivores) 赤鼻棱鳀(Thryssa chefuensis)、青鳞小沙丁(Sardinella zunasi)、鳀 (Engraulis japonicus)等 2 大泷六线鱼(Hexagrammos otakii) 大泷六线鱼(Hexagrammos otakii) 3 小眼绿鳍鱼(Chelidonichthys kumu) 小眼绿鳍鱼(Chelidonichthys kumu) 4 其他虾食性鱼类(Other shrimp predators) 矛尾鰕虎鱼(Chaeturichthys stigmatias)、皮氏叫姑鱼(Johnius belangerii)等 5 棘头梅童鱼(Collichthys lucidus) 棘头梅童鱼(Collichthys lucidus) 6 小黄鱼(Pseudosciaena polyactis) 小黄鱼(Pseudosciaena polyactis) 7 其他虾/鱼食性鱼类
(Other shrimp/fish predators)星康吉鳗(Conger myriaster)、白姑鱼(Argyrosomus argentatus)等 8 鱼食性鱼类(Piscivores) 长蛇鲻(Saurida elongata)、黄鮟鱇(Lophius litulon)、
带鱼(Trichiurus haumela)等9 方氏云鳚(Enedrias fangi) 方氏云鳚(Enedrias fangi) 10 其他底栖动物食性鱼类
(Other benthivores)六丝钝尾鰕虎鱼(Amblychaeturichthys hexanema)、角木叶鲽(Pleuronichthys cornutus)、
长丝鰕虎鱼(Cryptocentrus filifer)等11 口虾蛄(Oratosquilla oratoria) 口虾蛄(Oratosquilla oratoria) 12 戴氏赤虾(Metapenaeopsis dalei) 戴氏赤虾(Metapenaeopsis dalei) 13 其他虾类(Other Shrimps) 脊腹褐虾(Crangon affinis)、鹰爪虾(Trachysalambria curvirostris)、日本鼓虾(Alpheus japonicus)等 14 三疣梭子蟹(Portunus trituberculatus) 三疣梭子蟹(Portunus trituberculatus) 15 其他蟹类(Other Crabs) 日本蟳(Charybdis japonica)、双斑蟳(Charybdis bimaculata)、强壮菱蟹(Enoploambrus valida)等 16 枪乌贼 (Loligo sp.) 枪乌贼 (Loligo sp.) 17 大型头足类(Large cephalopoda) 长蛸(Octopus variabilis)、短蛸(Octopus ochellatus)、金乌贼(Sepia esculenta) 18 小型头足类(Small cephalopoda) 双喙耳乌贼(Sepiola birostrata)、四盘耳乌贼(Euprymna morsei) 19 其他软体动物(Other Molluscs) 双壳类(Bivalvia)、腹足类(Gastropoda) 20 多毛类(Polychates) 多毛类(Polychates) 21 端足类(Amphipods) 端足类(Amphipods) 22 棘皮动物(Echinoderms) 棘皮动物(Echinoderms) 23 其他底栖动物
(Other demersal invertebrate)日本浪飘水虱(Cirolana japonensis)等 24 浮游动物(Zooplankton) 中华哲水蚤(Calanus sinicus)、强壮箭虫(Sagitta crassa)等 25 浮游植物(Phytoplankton) 硅藻(Bacillariophyta)、甲藻(Pyrrophyta)等 26 碎屑(Detritus) 碎屑(Detritus) 表 2 海州湾及邻近海域生态系统Ecopath模型的基本参数
Tab. 2 Basic input data and estimated parameters for the Haizhou Bay and adjacent waters Ecopath model
编号 功能群 营养级 生物量/t·km−2 生产量/生物量(P/B) 消费量/生物量(Q/B) 生态营养效率(EE) 1 浮游动物食性鱼类 3.23 0.111 2.370 5.980 0.923 2 大泷六线鱼 3.72 0.025 2.900 9.000 0.468 3 小眼绿鳍鱼 3.98 0.248 1.479 4.230 0.449 4 其他虾食性鱼类 3.90 0.034 0.625 6.085 0.413 5 棘头梅童鱼 3.51 0.079 4.600 6.060 0.310 6 小黄鱼 3.84 0.164 1.660 5.900 0.837 7 其他虾/鱼食性鱼类 3.80 0.021 4.600 7.600 0.390 8 鱼食性鱼类 4.19 0.094 0.800 4.500 0.595 9 方氏云鳚 3.22 0.096 2.790 6.970 0.190 10 其他底栖动物食性鱼类 3.31 0.085 0.958 4.930 0.641 11 口虾蛄 3.37 0.057 1.340 7.430 0.956 12 戴氏赤虾 2.98 0.060 5.650 26.900 0.934 13 其他虾类 2.93 0.273 8.000 28.000 0.950 14 三疣梭子蟹 2.87 0.876 3.500 11.000 0.122 15 其他蟹类 2.85 0.039 3.500 12.000 0.931 16 枪乌贼 3.62 0.053 3.000 9.750 0.451 17 大型头足类 3.72 0.034 2.000 7.000 0.476 18 小型头足类 3.28 0.046 3.000 9.750 0.950 19 其他软体动物 2.18 5.200 6.000 27.000 0.251 20 多毛类 2.16 3.080 6.750 22.500 0.198 21 端足类 2.33 0.059 8.000 30.000 0.950 22 棘皮动物 2.51 3.080 1.200 3.580 0.623 23 其他底栖动物 2.18 1.967 1.570 8.600 0.950 24 浮游动物 2.05 2.271 25.000 122.100 0.989 25 浮游植物 1.00 20.673 106.520 0.141 26 碎屑 1.00 43.000 0.075 注:加粗数据表示估计参数。 表 3 海州湾及邻近海域生态系统内主要营养效应(前10位)
Tab. 3 The top ten trophic effects of the Haizhou Bay and adjacent waters ecosystems
效应 功能群 数值 下行效应 方氏云鳚 0.982 其他底栖动物食性鱼类 0.956 大型头足类 0.949 其他虾/鱼食性鱼类 0.919 鱼食性鱼类 0.905 口虾蛄 0.880 浮游动物食性鱼类 0.871 小眼绿鳍鱼 0.819 大泷六线鱼 0.818 小黄鱼 0.772 上行效应 浮游植物 0.967 端足类 0.949 碎屑 0.925 其他底栖动物 0.825 多毛类 0.752 其他蟹类 0.724 其他软体动物 0.666 小型头足类 0.648 棘头梅童类 0.476 其他虾类 0.410 表 4 海州湾及邻近海域生态系统的总体特征参数
Tab. 4 General characteristic parameters for the Haizhou Bay and adjacent waters ecosystem
参数 数值 总消耗量/t·km−2·a−1 542.975 总输出量/t·km−2·a−1 1 891.732 总呼吸量/t·km−2·a−1 310.309 流入碎屑总量/t·km−2·a−1 2 045.675 系统总流量/t·km−2·a−1 4 790.691 总生产量/t·km−2·a−1 2 326.112 总初级生产量/t·km−2·a−1 2 202.041 总初级生产量/总呼吸(TPP/TR) 7.096 净生产量/t·km−2·a−1 1 891.732 总初级生产量/总生物量(TPP/B) 56.866 总生物量(不计碎屑)/t·km−2 38.724 连接指数(CI) 0.429 系统杂食指数(SOI) 0.204 循环指数(FCI)/% 1.392 总能量转换效率/% 12.63 -
[1] 章守宇, 张焕君, 焦俊鹏, 等. 海州湾人工鱼礁海域生态环境的变化[J]. 水产学报, 2006, 30(4): 475−480.Zhang Shouyu, Zhang Huanjun, Jiao Junpeng, et al. Change of ecological environment of artificial reef waters in Haizhou Bay[J]. Journal of Fisheries of China, 2006, 30(4): 475−480. [2] 王冠钰, 郭佩芳. 基于生态系统的渔业管理方式(EBFM)在我国的适用性[J]. 海洋环境科学, 2014, 33(5): 792−797.Wang Guanyu, Guo Peifang. The adaptability of ecosystem-based fisheries management (EBFM) in China[J]. Marine Environmental Science, 2014, 33(5): 792−797. [3] Christensen V, Walters C J, Pauly D. Ecopath with Ecosim: a user’s guide[R]. Penang, Malaysia: Fisheries Centre, University of British Columbia, 2005. [4] Christensen V, Walters C J. Ecopath with Ecosim: methods, capabilities and limitations[J]. Ecological Modelling, 2004, 172(2/4): 109−139. [5] Polovina J J. Model of a coral reef ecosystem: I. The ECOPATH model and its application to French Frigate Shoals[J]. Coral Reefs, 1984, 3(1): 1−11. doi: 10.1007/BF00306135 [6] Ulanowicz R E. Growth and Development: Ecosystem Phenomenology[M]. New York: Springer Verlag, 1986. [7] Pauly D, Christensen V, Walters C. Ecopath, Ecosim, and Ecospace as tools for evaluating ecosystem impact of fisheries[J]. ICES Journal of Marine Science, 2000, 57(3): 697−706. doi: 10.1006/jmsc.2000.0726 [8] Link J, Col L, Guida V, et al. Response of balanced network models to large-scale perturbation: Implications for evaluating the role of small pelagics in the Gulf of Maine[J]. Ecological Modelling, 2009, 220(3): 351−369. doi: 10.1016/j.ecolmodel.2008.10.009 [9] Guo C B, Ye S W, Lek S, et al. The need for improved fishery management in a shallow macrophytic lake in the Yangtze River basin: evidence from the food web structure and ecosystem analysis[J]. Ecological Modelling, 2013, 267: 138−147. doi: 10.1016/j.ecolmodel.2013.07.013 [10] Han D Y, Xue Y, Zhang C L, et al. A mass balanced model of trophic structure and energy flows of a semi-closed marine ecosystem[J]. Acta Oceanologica Sinica, 2017, 36(10): 60−69. doi: 10.1007/s13131-017-1071-6 [11] 袁健美, 张虎, 贲成恺, 等. 海洲湾大型底栖动物群落组成及次级生产力[J]. 海洋渔业, 2018, 40(1): 19−26. doi: 10.3969/j.issn.1004-2490.2018.01.003Yuan Jianmei, Zhang Hu, Ben Chengkai, et al. Macrobenthic community composition and it’s secondary productivity in the Haizhou Bay[J]. Marine Fisheries, 2018, 40(1): 19−26. doi: 10.3969/j.issn.1004-2490.2018.01.003 [12] Tong Ling, Tang Qisheng, Pauly D. A preliminary approach on mass-balance ecopath model of the Bohai Sea[J]. Chinese Journal of Applied Ecology, 2000, 11(3): 435−440. [13] 林群, 王俊, 李忠义, 等. 黄河口邻近海域生态系统能量流动与三疣梭子蟹增殖容量估算[J]. 应用生态学报, 2015, 26(11): 3523−3531.Lin Qun, Wang Jun, Li Zhongyi, et al. Assessment of ecosystem energy flow and carrying capacity of swimming crab enhancement in the Yellow River Estuary and adjacent waters[J]. Chinese Journal of Applied Ecology, 2015, 26(11): 3523−3531. [14] 王腾, 张贺, 张虎, 等. 基于营养通道模型的海州湾中国明对虾生态容纳量[J]. 中国水产科学, 2016, 23(4): 965−975.Wang Teng, Zhang He, Zhang Hu, et al. Ecological carrying capacity of Chinese shrimp stock enhancement in Haizhou Bay of East China based on Ecopath model[J]. Journal of Fishery Sciences of China, 2016, 23(4): 965−975. [15] 张硕, 王腾, 符小明, 等. 连云港海州湾渔业生态修复水域生态系统能量流动模型初探[J]. 海洋环境科学, 2015, 34(1): 42−47.Zhang Shuo, Wang Teng, Fu Xiaoming, et al. A primary study on the energy flow in the ecosystem of fishery ecological restoration area in Haizhou Bay, Lianyungang[J]. Marine Environmental Science, 2015, 34(1): 42−47. [16] 欧阳力剑, 郭学武. 东、黄海主要鱼类Q/B值与种群摄食量研究[J]. 渔业科学进展, 2010, 31(2): 23−29. doi: 10.3969/j.issn.1000-7075.2010.02.004Ouyang Lijian, Guo Xuewu. Studies on the Q/B values and food consumption of major fishes in the East China Sea and the Yellow Sea[J]. Progress in Fishery Sciences, 2010, 31(2): 23−29. doi: 10.3969/j.issn.1000-7075.2010.02.004 [17] 徐超, 王思凯, 赵峰, 等. 基于Ecopath模型的长江口生态系统营养结构和能量流动研究[J]. 海洋渔业, 2018, 40(3): 309−318. doi: 10.3969/j.issn.1004-2490.2018.03.006Xu Chao, Wang Sikai, Zhao Feng, et al. Trophic structure and energy flow of the Yangtze Estuary ecosystem based on the analysis with Ecopath model[J]. Marine Fisheries, 2018, 40(3): 309−318. doi: 10.3969/j.issn.1004-2490.2018.03.006 [18] 隋昊志, 薛莹, 任一平, 等. 海州湾鱼类生态类群的研究[J]. 中国海洋大学学报, 2017, 47(12): 59−71.Sui Haozhi, Xue Ying, Ren Yiping, et al. Studies on the ecological groups of fish communities in Haizhou Bay, China[J]. Periodical of Ocean University of China, 2017, 47(12): 59−71. [19] 邓景耀, 孟田湘, 任胜民. 渤海鱼类食物关系的初步研究[J]. 生态学报, 1986, 6(4): 356−364.Deng Jingyao, Meng Tianxiang, Ren Shengmin. Food web of fishes in Bohai Sea[J]. Acta Ecologica Sinica, 1986, 6(4): 356−364. [20] 程济生, 朱金声. 黄海主要经济无脊椎动物摄食特征及其营养层次的研究[J]. 海洋学报, 1997, 19(6): 102−108.Cheng Jisheng, Zhu Jinsheng. Study on feeding characteristics and nutrient level of main economic invertebrates in the Yellow Sea[J]. Haiyang Xuebao, 1997, 19(6): 102−108. [21] 杨纪明. 渤海鱼类的食性和营养级研究[J]. 现代渔业信息, 2001, 16(10): 10−19.Yang Jiming. A study on food and trophic levels of Baohai Sea fish[J]. Modern Fisheries Information, 2001, 16(10): 10−19. [22] 张波. 东、黄海带鱼的摄食习性及随发育的变化[J]. 海洋水产研究, 2004, 25(2): 6−12.Zhang Bo. Feeding habits and ontogenetic diet shift of hairtail fish (Trichiurus lepturus) in East China Sea and Yellow Sea[J]. Marine Fisheries Research, 2004, 25(2): 6−12. [23] 徐开达, 金海卫, 卢占晖, 等. 东海区短鳄齿鱼摄食生态的初步研究[J]. 海洋科学, 2012, 36(7): 79−88.Xu Kaida, Jin Haiwei, Lu Zhanhui, et al. Preliminary study on feeding ecology of Champsodon snyderi in East China Sea region[J]. Marine Sciences, 2012, 36(7): 79−88. [24] 杨纪明, 谭雪静. 渤海3种头足类食性分析[J]. 海洋科学, 2000, 24(4): 53−55. doi: 10.3969/j.issn.1000-3096.2000.04.017Yang Jiming, Tan Xuejing. Food analysis of three cephalopod species in the Bohai Sea[J]. Marine Sciences, 2000, 24(4): 53−55. doi: 10.3969/j.issn.1000-3096.2000.04.017 [25] 杨纪明. 渤海涟虫类和软体动物幼虫食性的观察[J]. 海洋科学, 1998(6): 36−38.Yang Jiming. Observations on food of cumaceans and post larvae of mollusks in the Bohai Sea[J]. Marine Sciences, 1998(6): 36−38. [26] 杨德渐, 孙世春, 宋微波, 等. 海洋无脊椎动物学[M]. 青岛: 中国海洋大学出版社, 1999.Yang Dejian, Sun Shichun, Song Weibo, et al. Marine Invertebrate[M]. Qingdao: China Ocean University Press, 1999. [27] 邓景耀, 赵传铟. 海洋渔业生物学[M]. 北京: 农业出版社, 1991.Deng Jingyao, Zhao Chuanyin. Marine Fishery Biology[M]. Beijing: China Agriculture Press, 1991. [28] 赵文. 水生生物学[M]. 北京: 中国农业出版社, 2005.Zhao Wen. Hydrobiology[M]. Beijing: China Agriculture Press, 2005. [29] Leontief W W. The structure of the U.S. economy[J]. Scientific American, 1965, 212(4): 25−35. doi: 10.1038/scientificamerican0465-25 [30] Ulanowicz R E, Puccia C J. Mixed trophic impacts in ecosystems[J]. Coenoses, 1990, 5(1): 7−16. [31] Libralato S, Christensen V, Pauly D. A method for identifying keystone species in food web models[J]. Ecological Modelling, 2006, 195(3/4): 153−171. [32] Morissette L. Complexity, cost and quality of ecosystem models and their impact on resilience: a comparative analysis, with emphasis on marine mammals and the Gulf of St. Lawrence[D]. Vancouver: University of British Columbia, 2007. [33] 任晓明, 徐宾铎, 张崇良, 等. 海州湾及邻近海域鱼类群落的营养功能群及其动态变化[J]. 中国水产科学, 2019, 26(1): 141−150. doi: 10.3724/SP.J.1118.2019.18149Ren Xiaoming, Xu Binduo, Zhang Chongliang, et al. The composition of and variations in the trophic guilds of fish assemblages in Haizhou Bay and adjacent waters[J]. Journal of Fishery Sciences of China, 2019, 26(1): 141−150. doi: 10.3724/SP.J.1118.2019.18149 [34] Odum E P. The strategy of ecosystem development[J]. Science, 1969, 164(3877): 262−270. doi: 10.1126/science.164.3877.262 [35] 林群, 王俊, 李忠义, 等. 黄河口邻近水域贝类生态容量[J]. 应用生态学报, 2018, 29(9): 3131−3138.Lin Qun, Wang Jun, Li Zhongyi, et al. Ecological carrying capacity of shellfish in the Yellow River Estuary and its adjacent waters[J]. Chinese Journal of Applied Ecology, 2018, 29(9): 3131−3138. [36] 王在峰. 海州湾海洋特别保护区生态恢复适宜性评估[D]. 南京: 南京师范大学, 2011.Wang Zaifeng. Research on ecological recovery suitability assessment for Haizhou Bay special marine[D]. Nanjing: Nanjing Normal University, 2011. [37] Bitterlich G, Gnaiger E. Phytoplanktivorous or omnivorous fish? Digestibility of zooplankton by silvercarp, Hypophthalmichthys molitrix (Val.)[J]. Aquaculture, 1984, 40(3): 261−263. doi: 10.1016/0044-8486(84)90194-7 [38] Michener R M, Kaufman L. Stable isotope ratios as tracers in marine aquatic food webs[M]//Michener R M, Lajtha K. Stable Isotopes in Ecology and Environmental Science. Oxford: Blackwell Publishing, 1994: 138−186. [39] Gu B, Schell D M, Huang X, et al. Stable isotope evidence for dietary overlap between two planktivorous fishes in aquaculture ponds[J]. Canadian Journal of Fisheries and Aquatic Sciences, 1996, 53(12): 2814−2818. doi: 10.1139/f96-242 [40] Fry B. Stable Isotope Ecology[M]. New York: Springer, 2006. [41] Cerling T E, Ehleringer J R, Harris J M. Carbon dioxide starvation, the development of C4 ecosystems, and mammalian evolution[J]. Philosophical Transactions of the Royal Society B: Biological Sciences, 1998, 353(1365): 159−171. doi: 10.1098/rstb.1998.0198