鱼类自然死亡系数评估研究进展
|
摘要
自然死亡系数是渔业资源评估中不可或缺的重要参数,其准确度直接决定了资源评估结果的可靠性,进而影响渔业管理策略的制定。本文从生活史参数、标志回捕和年龄结构3个方面列举了国内外自然死亡系数的常用评估方法,讨论了相关方法的优缺点及影响因素,并以犬齿牙鲆(Paralichthys dentatus)和中国近海鱼类为例对比分析不同模型的计算结果。在此基础上,着重介绍了Pauly经验公式在中国近海主要经济鱼类自然死亡系数评估中的应用进展及存在问题。根据渔业资源调查和研究数据现状,认为现阶段使用Pauly经验公式评估中国近海经济鱼类自然死亡系数具有积极作用。
Natural mortality coefficient(M)is a key parameter in fisheries stock assessment.Its accuracy directly determines the reliability of assessment results and also influences the establishment of fisheries management.Most mathematical models of fish stock dynamics incorporate M,either directly or indirectly.Fishery scientists have explored accurate estimates of M for a long time,in order to develop a more precise understanding of fish populations and to establish a more scientific basis for managing fisheries resources and sustainable development.The present study introduces and evaluates the most common models for estimating natural mortality coefficients for fish populations from three aspects,namely life-history parameters,tag-return,and age-structure.The first approach to estimate the instantaneous rate of natural mortality in fish stocks is based on the observation that M is strongly correlated with life history parameters,such as maximum observed age,age at reproductive maturity,asymptotic fish length,and growth rate.The Pauly empirical equation,Gislason empirical equation,and Tanaka’s maximum observed age’s method are among the most commonly used methods for assessing inshore fisheries in China.By classifying tagging data,based on catch per unit effort,size at release,and the probability of tag loss,Chapman,Hampton,and Treble have each constructed their own models,and some scholars have combined these methods to develop a model that is superior to any of the individual models.The assessment results of using tag-return methods would have a high reliability,but the huge costs of investigation,lower credibility of recapture,and prior model assumptions contribute to poor generality.Considering that age is the potentially most important factor affecting estimates of M,virtual population analysis(VPA)and cohort analysis(CA)are established using catch-at-age data.These methods could be used to reliably estimate fish stocks with long-term catch data.However,considering the incomplete and inaccurate statues of historical catch data in Chinese coastal fishes,the practicality of the VPA or CA models is relatively low.Meanwhile,taking Paralichthys dentatus and some coastal fish stocks as examples,this study compared estimation results through different methods and found that Pauly empirical equation is more reasonable to calculate M.Then,this paper introduced the misapplications of Pauly empirical equation in inshore fisheries of China.The main misapplications were equation error,unit error and calculation mistake.According to the limited data and fishery exploitation status,we suggest that Pauly empirical equation has a positive meaning to assess the M for major economic fish stocks in China.
Natural mortality coefficient(M)is a key parameter in fisheries stock assessment.Its accuracy directly determines the reliability of assessment results and also influences the establishment of fisheries management.Most mathematical models of fish stock dynamics incorporate M,either directly or indirectly.Fishery scientists have explored accurate estimates of M for a long time,in order to develop a more precise understanding of fish populations and to establish a more scientific basis for managing fisheries resources and sustainable development.The present study introduces and evaluates the most common models for estimating natural mortality coefficients for fish populations from three aspects,namely life-history parameters,tag-return,and age-structure.The first approach to estimate the instantaneous rate of natural mortality in fish stocks is based on the observation that M is strongly correlated with life history parameters,such as maximum observed age,age at reproductive maturity,asymptotic fish length,and growth rate.The Pauly empirical equation,Gislason empirical equation,and Tanaka’s maximum observed age’s method are among the most commonly used methods for assessing inshore fisheries in China.By classifying tagging data,based on catch per unit effort,size at release,and the probability of tag loss,Chapman,Hampton,and Treble have each constructed their own models,and some scholars have combined these methods to develop a model that is superior to any of the individual models.The assessment results of using tag-return methods would have a high reliability,but the huge costs of investigation,lower credibility of recapture,and prior model assumptions contribute to poor generality.Considering that age is the potentially most important factor affecting estimates of M,virtual population analysis(VPA)and cohort analysis(CA)are established using catch-at-age data.These methods could be used to reliably estimate fish stocks with long-term catch data.However,considering the incomplete and inaccurate statues of historical catch data in Chinese coastal fishes,the practicality of the VPA or CA models is relatively low.Meanwhile,taking Paralichthys dentatus and some coastal fish stocks as examples,this study compared estimation results through different methods and found that Pauly empirical equation is more reasonable to calculate M.Then,this paper introduced the misapplications of Pauly empirical equation in inshore fisheries of China.The main misapplications were equation error,unit error and calculation mistake.According to the limited data and fishery exploitation status,we suggest that Pauly empirical equation has a positive meaning to assess the M for major economic fish stocks in China.
引文
[1]Lee H H,Maunder M N,Piner K R,et al.Estimating natural mortality within a fisheries stock assessment model:An evaluation using simulation analysis based on twelve stock assessments[J].Fisheries Research,2011,109(1):89-94.
[2]Sparre P,Ursin E,Venema S C,et al.Introduction to Tropical Fish Stock Assessment[M].Rome:Food and Agriculture Organization of the United Nations,1998:117-118.
[3]Schaefer M B.Some aspects of the dynamics of populations important to the management of the commercial marine fisheries[J].Inter-American Tropical Tuna Commission Bulletin,1954,1(2):23-56.
[4]Pella J J,Tomlinson P K.A generalized stock production model[J].Inter-American Tropical Tuna Commission Bulletin,1969,13(3):419-497.
[5]Aanes S,Engen S,S?ther B E,et al.Estimation of the parameters of fish stock dynamics from catch-at-age data and indices of abundance:can natural and fishing mortality be separated?[J].Canadian Journal of Fisheries and Aquatic Sciences,2007,64(8):1130-1142.
[6]Gislason H,Daan N,Rice J C,et al.Size,growth,temperature and the natural mortality of marine fish[J].Fish and Fisheries,2010,11(2):149-158.
[7]Clark W G.Effects of an erroneous natural mortality rate on a simple age-structured stock assessment[J].Canadian Journal of Fisheries and Aquatic Sciences,1999,56(10):1721-1731.
[8]Zhan B Y.Fish Stock Assessment[M].Beijing:China Agriculture Press,1995:109-113.[詹秉义.渔业资源评估[M].北京:中国农业出版社,1995:109-113.]
[9]Hamel O S.A method for calculating a meta-analytical prior for the natural mortality rate using multiple life history correlates[J].ICES Journal of Marine Science,2014,72(1):62-69.
[10]Zheng J.Uncertainties of natural mortality estimates for eastern Bering Sea snow crab,Chionoecetes opilio[J].Fisheries Research,2003,65(1-3):411-425.
[11]Hewitt D A,Hoenig J M.Comparison of two approaches for estimating natural mortality based on longevity[J].Fishery Bulletin,2005,103(2):433-437.
[12]Vetter E.Estimation of natural mortality in fish stocks:a review[J].Fishery Bulletin,1988,86(1):25-43.
[13]Wang Y B.Estimation of fish natural mortality coefficient using virtual population analysis[D].Qingdao:Ocean University of China,2007:12-18.[王迎宾.应用实际种群分析(VPA)求解鱼类自然死亡系数的研究[D].青岛:中国海洋大学,2007:12-18.]
[14]Pauly D.On the interrelationships between natural mortality,growth parameters,and mean environmental temperature in175 fish stocks[J].ICES Journal of Marine Science,1980,39(2):175-192.
[15]Sippel T,Lee H H,Piner K,et al.Searching for M:Is there more information about natural mortality in stock assessments than we realize?[J].Fisheries Research,2017,192:135-140.
[16]Zhang Y X,Miao Z Q.Review of fish assessment models[J].Journal of Zhejiang Ocean University(Natural Science),2006,25(3):305-311.[张月霞,苗振清.渔业资源的评估方法和模型研究进展[J].浙江海洋学院学报(自然科学版),2006,25(3):305-311.]
[17]Quinn T J,Deriso R B.Quantitative Fish Dynamics[M].Oxford:Oxford University Press,1999.
[18]Jensen A.Beverton and Holt life history invariants result from optimal trade-off of reproduction and survival[J].Canadian Journal of Fisheries and Aquatic Sciences,1996,53(4):820-822.
[19]Lorenzen K.The relationship between body weight and natural mortality in juvenile and adult fish:a comparison of natural ecosystems and aquaculture[J].Journal of Fish Biology,1996,49(4):627-642.
[20]Peterson I,Wroblewski J.Mortality rate of fishes in the pelagic ecosystem[J].Canadian Journal of Fisheries and Aquatic Sciences,1984,41(7):1117-1120.
[21]Post J R,Evans D O.Size-dependent overwinter mortality of young-of-the-year yellow perch(Perca flavescens):laboratory,in situ enclosure,and field experiments[J].Canadian Journal of Fisheries and Aquatic Sciences,1989,46(11):1958-1968.
[22]Mc Gurk M D.Natural mortality of marine pelagic fish eggs and larvae:role of spatial patchiness[J].Marine Ecology Progress Series,1986,34(3):227-242.
[23]Pascual M A,Iribarne O O.How good are empirical predictions of natural mortality?[J].Fisheries Research,1993,16(1):17-24.
[24]Beverton R J H,Holt S J.The Lifespan of Animals(Colloquia on Ageing):A review of the lifespans and mortality rates of fishes in nature,and their relation to growth and other physiological characteristics[M].CIBA Foundation Symposium,1959,5:142-179.
[25]Kenchington T J.Natural mortality estimators for information-limited fisheries[J].Fish and Fisheries,2014,15(4):533-562.
[26]Venema S C,Christensen J,Pauly D.Contributions to tropical fisheries biology.Papers prepared by the participants at the FAO/DANIDA follow-up training courses on fish stock assessment in the Tropics,Hirtshals,Denmark 5-30 May1986,Manila,Philippines 12 January-6 February 1987[R].FAO Fisheries Report,389:519.
[27]Gislason H,Daan N,Rice J,et al.Does natural mortality depend on individual size?[C].ICES Science Conference,2008/F:16.
[28]Xu H L,Chen Y,Chen X J,et.al.Comparison of two empirical estimates of natural mortality for Chinese shrimp[J].Joural of Fiseriers of China,2016,40(1):45-54.[徐海龙,陈勇,陈新军,等.两种经验方法估算中国明对虾自然死亡的比较[J].水产学报,2016,40(1):45-54.]
[29]Tanaka S.Studies on the dynamics and management of fish populations[J].Bulletin of the Tokai Regional Fisheries Research Laboratory,1960,28:1-200.
[30]Zhan B Y,Lou D C,Zhong J S.An assessment of the filefish population and rational exploitation of the resource[J].Joural of Fiseriers of China,1986,10(4):409-418.[詹秉义,楼冬春,钟俊生.绿鳍马面鲀资源评析与合理利用[J].水产学报,1986,10(4):409-418.]
[31]Alverson D L,Carney M J.A graphic review of the growth and decay of population cohorts[J].ICES Journal of Marine Science,1975,36(2):133-143.
[32]Montgomery S,Barchia I,Walsh C.Estimating rates of mortality in stocks of Metapenaeus macleayi in estuaries of eastern Australia[J].Fisheries Research,2012,113(1):55-67.
[33]Gunderson D R,Zimmermann M,Nichol D G,et al.Indirect estimates of natural mortality rate for arrowtooth flounder(Atheresthes stomias)and darkblotched rockfish(Sebastes crameri)[J].Fishery Bulletin,2003,101(1):175-182.
[34]Pollock K H,Hoenig J M,Jones C M.Estimation of fishing and natural mortality when a tagging study is combined with a creel survey or port sampling[J].American Fisheries Society Symposium,1991,12:423-434.
[35]Williams B K,Nichols J D,Conroy M J.Analysis and Management of Animal Populations[M].San Diego:Academic Press,2002.
[36]Chapman F A,Hartless C S,Carr S H.Population size estimates of sturgeon in the Suwannee River,Florida,U.S.A[J].Gulf of Mexico Science,1997,2:88-91.
[37]Hilborn R,Walters C J.Quantitative Fisheries Stock Assessment:Choice,Dynamics and Uncertainty[M].New York:Chapman and Hall,1992.
[38]Hampton J.Natural mortality rates in tropical tunas:size really does matter[J].Canadian Journal of Fisheries and Aquatic Sciences,2000,57(5):1002-1010.
[39]Treble R J,Day R W,QuinnⅡT J.Detection and effects on mortality estimates of changes in tag loss[J].Canadian Journal of Aquatic Science,1993,50:1435-1441.
[40]Wetherall J A.Analysis of double tagging experiments[J].Fishery Bulletin,1982,80(4):687-701.
[41]Brooks E N,Pollock K H,Hoenig J M,et al.Estimation of fishing and natural mortality from tagging studies on fisheries with two user groups[J].Canadian Journal of Fisheries and Aquatic Sciences,1998,55(9):2001-2010.
[42]Pine W E,Pollock K H,Hightower J E,et al.A review of tagging methods for estimating fish population size and components of mortality[J].Fisheries Research,2003,28(10):10-23.
[43]Kenneth H P,Honghua J,Joseph E H.Combining telemetry and fisheries tagging models to estimate fishing and natural mortality rates[J].Transactions of the American Fisheries Society,2004,133(3):639-648.
[44]Thorley J L,Andrusak G F.The fishing and natural mortality of large,piscivorous Bull Trout and Rainbow Trout in Kootenay Lake,British Columbia(2008-2013)[J].Peer J,2017,5:e2874.
[45]Megrey B A.Review and comparison of age-structured stock assessment models from theoretical and applied points of view[J].American Fisheries Society Simposium,1989,6:8-48.
[46]Gulland J.Estimation of mortality rates.Annex to Arctic fisheries working group report[R].International Council for the Exploration of the Sea,C.M,1965 Doc.No.3,1965.
[47]Pope J.An investigation of the accuracy of virtual population analysis using cohort analysis[J].ICNAF Research Bulletin,1972,9(10):65-74.
[48]Guan W J,Tian S Q,Zhu J F,et al.A review of fisheries stock assessment models[J].Journal of Fishery Sciences of China,2013,20(5):1112-1120.[官文江,田思泉,朱江峰,等.渔业资源评估模型的研究现状与展望[J].中国水产科学,2013,20(5):1112-1120.]
[49]Maunder M N,Punt A E.A review of integrated analysis in fisheries stock assessment[J].Fisheries Research,2013,142(6):61-74.
[50]Liu Q,Wang Y J.Preliminary study of the tuning methods in fisheries virtual population analysis(VPA)[J].South China Fisheries Science,2007,3(1):1-6.[刘群,王艳君.对渔业实际种群分析(VPA)中调谐方法的初步研究[J].南方水产,2007,3(1):1-6.]
[51]Maunder M N,Wong R A.Approaches for estimating natural mortality:Application to summer flounder(Paralichthys dentatus)in the U.S.mid-Atlantic[J].Fisheries Research,2011,111(1):92-99.
[52]Gunderson D R.Trade-off between reproductive effort and adult survival in oviparous and viviparous fishes[J].Canadian Journal of Fisheries and Aquatic Sciences,1997,54(5):990-998.
[53]Froese R,Pauly D.“Fish Base,”World Wide Web electronic publication,Version(09/2010)[OL].http://www.fishbase.org/search.php.
[54]Hoenig J M.Empirical use of longevity data to estimate mortality rates[J].Fisheries Bulletin,1983,82(1):898-903
[55]Fei H N,Zhang S Q.Fishery Resources[M].Beijing:Science and Technology of China Press,1990:308-313.[费鸿年,张诗全.水产资源学[M].北京:中国科学技术出版社,1990:308-313.]
[56]Tang Q.Modification of the ricker stock recruitment model to account for environmentally induced variation in recruitment with particular reference to the blue crab fishery in Chesapeake Bay[J].Fisheries Research,1985,3(1):13-21.
[57]Sun D R,Qiu Y S.Esitimation of growth and mortality parameter of Priacanthus macracanthus in Beibu Bay[J].Journal of Fujian Fisheries,2003(1):7-12.[孙典荣,邱永松.北部湾短尾大眼鲷生长和死亡参数的估算[J].福建水产,2003(1):7-12.]
[58]Zhang K,Chen Z Z,Wang Y Z,et al.Population structure of Priacanthus macracanthus in the Beibu Gulf,and parameters for its growth,mortality and maturity[J].Journal of Tropical Oceanography,2016,35(5):20-28.[张魁,陈作志,王跃中,等.北部湾短尾大眼鲷群体结构及生长、死亡和性成熟参数估计[J].热带海洋学报,2016,35(5):20-28.]
[59]Sun D R,Qiu Y S.Estimation of growth and mortality parameters of Priacanthus macracanthus in the North Continental Shelf of South China Sea[J].Journal of Guangdong Ocean University,2004,24(1):28-34.[孙典荣,邱永松.南海北部大陆架短尾大眼鲷生长和死亡参数估计[J].广东海洋大学学报,2004,24(1):28-34.]
[60]Shu L M,Qiu Y S.Estimate for growth,mortality parameters and first capture specification suggestion of Saurida undosquamis(Richardson)in Northern South China Sea[J].Journal of Zhanjiang Ocean University,2004,24(3):29-35.[舒黎明,邱永松.南海北部花斑蛇鲻生长死亡参数估计及开捕规格[J].湛江海洋大学学报,2004,24(3):29-35.]
[61]Chen G B,Qiu Y S.Growth,mortality and rational utilization of horse Mackerel in Northern South China Sea[J].Journal of Zhanjiang Ocean University,2004,24(1):35-40.[陈国宝,邱永松.南海北部竹荚鱼的生长、死亡及合理利用[J].湛江海洋大学学报,2004,24(1):35-40.]
[62]Zhu J F,Qiu Y S.Growth and mortality of hairtails and their dynamic pool models in the northern South China Sea[J].Acta Oceanologica Sinica,2005,27(6):93-99.[朱江峰,邱永松.南海北部带鱼生长死亡与参数动态综合模式[J].海洋学报,2005,27(6):93-99.]
[63]Lin L S,Zheng Y J,Chen J H,et al.A preliminary study on fishery biology of main commercial fishes surveyed from the bottom trawl fisheries in the East China Sea[J].Marine Science,2006,30(2):21-25.[林龙山,郑元甲,程家骅,等.东海区底拖网渔业主要经济鱼类渔业生物学的初步研究[J].海洋科学,2006,30(2):21-25.]
[64]Zhang G Z,Li X S,Jin X S,et al.Changes of biological characteristics of small yellow croker(Larimichthys polyactis)in the central and southern Yellow Sea[J].Acta Ecological Sinica,2010,30(24):6854-6861.[张国政,李显森,金显仕,等.黄海中南部小黄鱼生物学特征的变化[J].生态学报,2010,30(24):6854-6861.]
[65]Zhang G Z,Li X S,Jin X S,et al.Growth,mortality and optimum catchable size of small yellow croaker(Larimichthys polyactis Bleeker)in the Southern Yellow Sea[J].Journal of Fishery Sciences of China,2010,17(4):839-846.[张国政,李显森,金显仕,等.黄海南部小黄鱼生长、死亡和最适开捕体长[J].中国水产科学,2010,17(4):839-846.]
[66]Liu Z L,Xie H Y,Yan L P,et al.Comparative population dynamics of small yellow croaker Larimichthys polyactis in Southern Yellow Sea and East China Sea[J].Journal of Dalian Ocean University,2013,28(6):627-632.[刘尊雷,谢汉阳,严利平,等.黄海南部和东海小黄鱼资源动态的比较[J].大连海洋大学学报,2013,28(6):627-632.]
[67]Liu Q,Xu B D,Ye Z J,et.al.Growth and Mortality of small yellow croaker(Larimichthys polyactis)inhabiting Haizhou Bay of China[J].Journal of Ocean University of China,2012,11(4):557-561.
[68]Wang Y B,Yu C G,Yu C D,et al.Preliminary research on the rational utilization of Psenopsis anomala in the offshore water of South Zhejiang[J].Journal of Zhejiang Ocean University(Natural Science),2010,29(2):114-119.[王迎宾,俞存根,虞聪达,等.浙江南部外海刺鲳资源合理利用的初步研究[J].浙江海洋学院学报(自然科学版),2010,29(2):114-119.]
[69]Du J G,Lu Z B,Chen M R,et al.Changes in ecological parameters and resources of Japanese butterfish,Psenopsis anomala,in the middle and north Taiwan Strait[J].Journal of Oceanography in Taiwan Strait,2010,29(2):234-240.[杜建国,卢振彬,陈明茹,等.台湾海峡中北部海域刺鲳种群生态学参数及其变动趋势[J].台湾海峡,2010,29(2):234-240.]
[70]Zhang K,Chen Z Z,Qiu Y S.Decadal changes in growth,mortality and maturity parameters of Evynnis cardinalis in Beibu Gulf[J].South China Fisheries Science,2016,12(6):9-16.[张魁,陈作志,邱永松.北部湾二长棘犁齿鲷生长、死亡和性成熟参数的年际变化[J].南方水产科学,2016,12(6):9-16.].
[71]Hou G,Zhu X L,Lu H S.Growth,mortality and population composition of crimson sea bream,Paragyrops edita Tanaka in Beibu Gulf[J].Journal of Guangdong Ocean University,2008,28(3):50-55.[侯刚,朱立新,卢伙胜.北部湾二长棘鲷生长、死亡及其群体组成[J].广东海洋大学学报,2008,28(3):50-55.]
[72]Chen Z Z,Qiu Y S.Esitimation of growth and mortality parameters of Parargyrops edita Tanaka in Beibu Bay[J].Journal of Fisheries of China,2003,27(3):251-257.[陈作志,邱永松.北部湾二长棘鲷生长和死亡参数估计[J].水产学报,2003,27(3):251-257.]
[73]Feng B,Hou G,Lu H S.Management recommendation for Paragyrops edita in Beibu Gulf based on per-recruitment analysis[J].System Science and Comprehensive Studies in Agriculture,2009,25(1):114-119.[冯波,侯刚,卢伙胜.北部湾二长棘鲷资源开发保护策略研究[J].农业系统科学与综合研究,2009,25(1):114-119.]
[74]Wang X H,Qiu Y S,Du F Y.Study on the growth,mortality and optimum catchable size of Nemipterus virgatus in the Northern South China Sea[J].Journal of Ocean University of China,2004,34(2):224-230.[王雪辉,邱永松,杜飞雁.南海北部金线鱼生长、死亡和最适开捕体长研究[J].中国海洋大学学报,2004,34(2):224-230.]
[75]Wang X H,Qiu Y S,Du F Y.Study on the growth,mortality and optimum catchable size of Nemipterus Virgatus in the Northern South China Sea[J].Journal of Ocean University of China,2004,34(2):224-230.[王雪辉,邱永松,杜飞雁.南海北部金线鱼生长、死亡和最适开捕体长研究[J].中国海洋大学学报,2004,34(2):224-230.]
[76]Lu Z B,Du J G.Growth and mortality character of Nemi pterus virgatus in southern Taiwan Strait[J].Journal of Tropical Ocean Nography,2008,27(5):75-79.[卢振彬,杜建国.台湾海峡南部金线鱼的生长与死亡特性[J].热带海洋学报,2008,27(5):75-79.]
[77]Liu Y W,Xue Y,Wei B F,et al.An estimation of population parameters for five major fish species in the Haizhou Bay,Yellow Sea[J].Journal of Fishery Sciences of China,2014,21(1):125-133.[刘元文,薛莹,魏邦福,等.海州湾5种主要鱼类种群参数估算[J].中国水产科学,2014,21(1):125-133.]
[78]Liu J D,Lu H S,Zhu L X,et al.The difference of growth,mortality and group composition Between female and male Saurida tumbil in the Beibu Gulf[J].Marine Fisheries,2009,31(3):243-253.[刘金殿,卢伙胜,朱立新,等.北部湾多齿蛇鲻雌雄群体组成、生长、死亡特征的差异[J].海洋渔业,2009,31(3):243-253.]
[79]Shu L M,Qiu Y S.Biology analysis of Saurida tumbil in northern South China Sea[J].Journal of Fishery Sciences of China,2004,11(2):154-158.[舒黎明,邱永松,南海北部多齿蛇鲻生物学分析[J].中国水产科学,2004,11(2):154-158.]
[80]Liu Y,Cheng J H,Li S F.Utilization status of Setipinna taty in the East China Sea and its rational exploitation[J].Journal of Fishery Sciences of China,2006,13(3):485-491.[刘勇,程家骅,李圣法.东海区黄鲫资源的利用现状及合理利用探讨[J].中国水产科学,2006,13(3):485-491.]
[81]Lu Z B,Chen X.The parameters and their changes of the growth and mortality of Engraulidae and Clupeidae in Fujian coastal waters[J].Journal of Xiamen University(Natural Science),2008,47(2):279-285.[卢振彬,陈骁.福建沿海几种鲱、鳀科鱼类生长与死亡参数及其变化[J].厦门大学学报(自然科学版),2008,47(2):279-285.]
[82]Chen G B,Qiu Y S.Study on growth,mortality and reasonable utilization of Decapterus maruadsi in northern continental shelf waters of South China Sea[J].Journal of Oceanography in Taiwan Strait,2003,22(4):457-464.[陈国宝,邱永松.南海北部陆架区蓝圆鲹的生长、死亡及合理利用研究[J].台湾海峡,2003,22(4):457-464.]
[83]Zheng J,Wang Y B,Li R X,et al.Stock assessment of Jack Scomber japonicus and Decapterus maruadsi in coastal waters of Zhejiang Province[J].Journal of Zhejiang Ocean University(Natural Science),2012,31(4):309-315.[郑基,王迎宾,李仁星,等.浙江海域鲐鲹鱼资源量评估[J].浙江海洋学院学报(自然科学版),2012,31(4):309-315.]
[84]Costello C,Ovando D,Hilborn R,et al.Status and Solutions for the World’s Unassessed Fisheries[J].Science,2012,338(6106):517-520.
[85]Chen Y.Quality of fisheries data and uncertainty in stock assessment[J].Scientia Marina,2003,67(S1):75-87.
[2]Sparre P,Ursin E,Venema S C,et al.Introduction to Tropical Fish Stock Assessment[M].Rome:Food and Agriculture Organization of the United Nations,1998:117-118.
[3]Schaefer M B.Some aspects of the dynamics of populations important to the management of the commercial marine fisheries[J].Inter-American Tropical Tuna Commission Bulletin,1954,1(2):23-56.
[4]Pella J J,Tomlinson P K.A generalized stock production model[J].Inter-American Tropical Tuna Commission Bulletin,1969,13(3):419-497.
[5]Aanes S,Engen S,S?ther B E,et al.Estimation of the parameters of fish stock dynamics from catch-at-age data and indices of abundance:can natural and fishing mortality be separated?[J].Canadian Journal of Fisheries and Aquatic Sciences,2007,64(8):1130-1142.
[6]Gislason H,Daan N,Rice J C,et al.Size,growth,temperature and the natural mortality of marine fish[J].Fish and Fisheries,2010,11(2):149-158.
[7]Clark W G.Effects of an erroneous natural mortality rate on a simple age-structured stock assessment[J].Canadian Journal of Fisheries and Aquatic Sciences,1999,56(10):1721-1731.
[8]Zhan B Y.Fish Stock Assessment[M].Beijing:China Agriculture Press,1995:109-113.[詹秉义.渔业资源评估[M].北京:中国农业出版社,1995:109-113.]
[9]Hamel O S.A method for calculating a meta-analytical prior for the natural mortality rate using multiple life history correlates[J].ICES Journal of Marine Science,2014,72(1):62-69.
[10]Zheng J.Uncertainties of natural mortality estimates for eastern Bering Sea snow crab,Chionoecetes opilio[J].Fisheries Research,2003,65(1-3):411-425.
[11]Hewitt D A,Hoenig J M.Comparison of two approaches for estimating natural mortality based on longevity[J].Fishery Bulletin,2005,103(2):433-437.
[12]Vetter E.Estimation of natural mortality in fish stocks:a review[J].Fishery Bulletin,1988,86(1):25-43.
[13]Wang Y B.Estimation of fish natural mortality coefficient using virtual population analysis[D].Qingdao:Ocean University of China,2007:12-18.[王迎宾.应用实际种群分析(VPA)求解鱼类自然死亡系数的研究[D].青岛:中国海洋大学,2007:12-18.]
[14]Pauly D.On the interrelationships between natural mortality,growth parameters,and mean environmental temperature in175 fish stocks[J].ICES Journal of Marine Science,1980,39(2):175-192.
[15]Sippel T,Lee H H,Piner K,et al.Searching for M:Is there more information about natural mortality in stock assessments than we realize?[J].Fisheries Research,2017,192:135-140.
[16]Zhang Y X,Miao Z Q.Review of fish assessment models[J].Journal of Zhejiang Ocean University(Natural Science),2006,25(3):305-311.[张月霞,苗振清.渔业资源的评估方法和模型研究进展[J].浙江海洋学院学报(自然科学版),2006,25(3):305-311.]
[17]Quinn T J,Deriso R B.Quantitative Fish Dynamics[M].Oxford:Oxford University Press,1999.
[18]Jensen A.Beverton and Holt life history invariants result from optimal trade-off of reproduction and survival[J].Canadian Journal of Fisheries and Aquatic Sciences,1996,53(4):820-822.
[19]Lorenzen K.The relationship between body weight and natural mortality in juvenile and adult fish:a comparison of natural ecosystems and aquaculture[J].Journal of Fish Biology,1996,49(4):627-642.
[20]Peterson I,Wroblewski J.Mortality rate of fishes in the pelagic ecosystem[J].Canadian Journal of Fisheries and Aquatic Sciences,1984,41(7):1117-1120.
[21]Post J R,Evans D O.Size-dependent overwinter mortality of young-of-the-year yellow perch(Perca flavescens):laboratory,in situ enclosure,and field experiments[J].Canadian Journal of Fisheries and Aquatic Sciences,1989,46(11):1958-1968.
[22]Mc Gurk M D.Natural mortality of marine pelagic fish eggs and larvae:role of spatial patchiness[J].Marine Ecology Progress Series,1986,34(3):227-242.
[23]Pascual M A,Iribarne O O.How good are empirical predictions of natural mortality?[J].Fisheries Research,1993,16(1):17-24.
[24]Beverton R J H,Holt S J.The Lifespan of Animals(Colloquia on Ageing):A review of the lifespans and mortality rates of fishes in nature,and their relation to growth and other physiological characteristics[M].CIBA Foundation Symposium,1959,5:142-179.
[25]Kenchington T J.Natural mortality estimators for information-limited fisheries[J].Fish and Fisheries,2014,15(4):533-562.
[26]Venema S C,Christensen J,Pauly D.Contributions to tropical fisheries biology.Papers prepared by the participants at the FAO/DANIDA follow-up training courses on fish stock assessment in the Tropics,Hirtshals,Denmark 5-30 May1986,Manila,Philippines 12 January-6 February 1987[R].FAO Fisheries Report,389:519.
[27]Gislason H,Daan N,Rice J,et al.Does natural mortality depend on individual size?[C].ICES Science Conference,2008/F:16.
[28]Xu H L,Chen Y,Chen X J,et.al.Comparison of two empirical estimates of natural mortality for Chinese shrimp[J].Joural of Fiseriers of China,2016,40(1):45-54.[徐海龙,陈勇,陈新军,等.两种经验方法估算中国明对虾自然死亡的比较[J].水产学报,2016,40(1):45-54.]
[29]Tanaka S.Studies on the dynamics and management of fish populations[J].Bulletin of the Tokai Regional Fisheries Research Laboratory,1960,28:1-200.
[30]Zhan B Y,Lou D C,Zhong J S.An assessment of the filefish population and rational exploitation of the resource[J].Joural of Fiseriers of China,1986,10(4):409-418.[詹秉义,楼冬春,钟俊生.绿鳍马面鲀资源评析与合理利用[J].水产学报,1986,10(4):409-418.]
[31]Alverson D L,Carney M J.A graphic review of the growth and decay of population cohorts[J].ICES Journal of Marine Science,1975,36(2):133-143.
[32]Montgomery S,Barchia I,Walsh C.Estimating rates of mortality in stocks of Metapenaeus macleayi in estuaries of eastern Australia[J].Fisheries Research,2012,113(1):55-67.
[33]Gunderson D R,Zimmermann M,Nichol D G,et al.Indirect estimates of natural mortality rate for arrowtooth flounder(Atheresthes stomias)and darkblotched rockfish(Sebastes crameri)[J].Fishery Bulletin,2003,101(1):175-182.
[34]Pollock K H,Hoenig J M,Jones C M.Estimation of fishing and natural mortality when a tagging study is combined with a creel survey or port sampling[J].American Fisheries Society Symposium,1991,12:423-434.
[35]Williams B K,Nichols J D,Conroy M J.Analysis and Management of Animal Populations[M].San Diego:Academic Press,2002.
[36]Chapman F A,Hartless C S,Carr S H.Population size estimates of sturgeon in the Suwannee River,Florida,U.S.A[J].Gulf of Mexico Science,1997,2:88-91.
[37]Hilborn R,Walters C J.Quantitative Fisheries Stock Assessment:Choice,Dynamics and Uncertainty[M].New York:Chapman and Hall,1992.
[38]Hampton J.Natural mortality rates in tropical tunas:size really does matter[J].Canadian Journal of Fisheries and Aquatic Sciences,2000,57(5):1002-1010.
[39]Treble R J,Day R W,QuinnⅡT J.Detection and effects on mortality estimates of changes in tag loss[J].Canadian Journal of Aquatic Science,1993,50:1435-1441.
[40]Wetherall J A.Analysis of double tagging experiments[J].Fishery Bulletin,1982,80(4):687-701.
[41]Brooks E N,Pollock K H,Hoenig J M,et al.Estimation of fishing and natural mortality from tagging studies on fisheries with two user groups[J].Canadian Journal of Fisheries and Aquatic Sciences,1998,55(9):2001-2010.
[42]Pine W E,Pollock K H,Hightower J E,et al.A review of tagging methods for estimating fish population size and components of mortality[J].Fisheries Research,2003,28(10):10-23.
[43]Kenneth H P,Honghua J,Joseph E H.Combining telemetry and fisheries tagging models to estimate fishing and natural mortality rates[J].Transactions of the American Fisheries Society,2004,133(3):639-648.
[44]Thorley J L,Andrusak G F.The fishing and natural mortality of large,piscivorous Bull Trout and Rainbow Trout in Kootenay Lake,British Columbia(2008-2013)[J].Peer J,2017,5:e2874.
[45]Megrey B A.Review and comparison of age-structured stock assessment models from theoretical and applied points of view[J].American Fisheries Society Simposium,1989,6:8-48.
[46]Gulland J.Estimation of mortality rates.Annex to Arctic fisheries working group report[R].International Council for the Exploration of the Sea,C.M,1965 Doc.No.3,1965.
[47]Pope J.An investigation of the accuracy of virtual population analysis using cohort analysis[J].ICNAF Research Bulletin,1972,9(10):65-74.
[48]Guan W J,Tian S Q,Zhu J F,et al.A review of fisheries stock assessment models[J].Journal of Fishery Sciences of China,2013,20(5):1112-1120.[官文江,田思泉,朱江峰,等.渔业资源评估模型的研究现状与展望[J].中国水产科学,2013,20(5):1112-1120.]
[49]Maunder M N,Punt A E.A review of integrated analysis in fisheries stock assessment[J].Fisheries Research,2013,142(6):61-74.
[50]Liu Q,Wang Y J.Preliminary study of the tuning methods in fisheries virtual population analysis(VPA)[J].South China Fisheries Science,2007,3(1):1-6.[刘群,王艳君.对渔业实际种群分析(VPA)中调谐方法的初步研究[J].南方水产,2007,3(1):1-6.]
[51]Maunder M N,Wong R A.Approaches for estimating natural mortality:Application to summer flounder(Paralichthys dentatus)in the U.S.mid-Atlantic[J].Fisheries Research,2011,111(1):92-99.
[52]Gunderson D R.Trade-off between reproductive effort and adult survival in oviparous and viviparous fishes[J].Canadian Journal of Fisheries and Aquatic Sciences,1997,54(5):990-998.
[53]Froese R,Pauly D.“Fish Base,”World Wide Web electronic publication,Version(09/2010)[OL].http://www.fishbase.org/search.php.
[54]Hoenig J M.Empirical use of longevity data to estimate mortality rates[J].Fisheries Bulletin,1983,82(1):898-903
[55]Fei H N,Zhang S Q.Fishery Resources[M].Beijing:Science and Technology of China Press,1990:308-313.[费鸿年,张诗全.水产资源学[M].北京:中国科学技术出版社,1990:308-313.]
[56]Tang Q.Modification of the ricker stock recruitment model to account for environmentally induced variation in recruitment with particular reference to the blue crab fishery in Chesapeake Bay[J].Fisheries Research,1985,3(1):13-21.
[57]Sun D R,Qiu Y S.Esitimation of growth and mortality parameter of Priacanthus macracanthus in Beibu Bay[J].Journal of Fujian Fisheries,2003(1):7-12.[孙典荣,邱永松.北部湾短尾大眼鲷生长和死亡参数的估算[J].福建水产,2003(1):7-12.]
[58]Zhang K,Chen Z Z,Wang Y Z,et al.Population structure of Priacanthus macracanthus in the Beibu Gulf,and parameters for its growth,mortality and maturity[J].Journal of Tropical Oceanography,2016,35(5):20-28.[张魁,陈作志,王跃中,等.北部湾短尾大眼鲷群体结构及生长、死亡和性成熟参数估计[J].热带海洋学报,2016,35(5):20-28.]
[59]Sun D R,Qiu Y S.Estimation of growth and mortality parameters of Priacanthus macracanthus in the North Continental Shelf of South China Sea[J].Journal of Guangdong Ocean University,2004,24(1):28-34.[孙典荣,邱永松.南海北部大陆架短尾大眼鲷生长和死亡参数估计[J].广东海洋大学学报,2004,24(1):28-34.]
[60]Shu L M,Qiu Y S.Estimate for growth,mortality parameters and first capture specification suggestion of Saurida undosquamis(Richardson)in Northern South China Sea[J].Journal of Zhanjiang Ocean University,2004,24(3):29-35.[舒黎明,邱永松.南海北部花斑蛇鲻生长死亡参数估计及开捕规格[J].湛江海洋大学学报,2004,24(3):29-35.]
[61]Chen G B,Qiu Y S.Growth,mortality and rational utilization of horse Mackerel in Northern South China Sea[J].Journal of Zhanjiang Ocean University,2004,24(1):35-40.[陈国宝,邱永松.南海北部竹荚鱼的生长、死亡及合理利用[J].湛江海洋大学学报,2004,24(1):35-40.]
[62]Zhu J F,Qiu Y S.Growth and mortality of hairtails and their dynamic pool models in the northern South China Sea[J].Acta Oceanologica Sinica,2005,27(6):93-99.[朱江峰,邱永松.南海北部带鱼生长死亡与参数动态综合模式[J].海洋学报,2005,27(6):93-99.]
[63]Lin L S,Zheng Y J,Chen J H,et al.A preliminary study on fishery biology of main commercial fishes surveyed from the bottom trawl fisheries in the East China Sea[J].Marine Science,2006,30(2):21-25.[林龙山,郑元甲,程家骅,等.东海区底拖网渔业主要经济鱼类渔业生物学的初步研究[J].海洋科学,2006,30(2):21-25.]
[64]Zhang G Z,Li X S,Jin X S,et al.Changes of biological characteristics of small yellow croker(Larimichthys polyactis)in the central and southern Yellow Sea[J].Acta Ecological Sinica,2010,30(24):6854-6861.[张国政,李显森,金显仕,等.黄海中南部小黄鱼生物学特征的变化[J].生态学报,2010,30(24):6854-6861.]
[65]Zhang G Z,Li X S,Jin X S,et al.Growth,mortality and optimum catchable size of small yellow croaker(Larimichthys polyactis Bleeker)in the Southern Yellow Sea[J].Journal of Fishery Sciences of China,2010,17(4):839-846.[张国政,李显森,金显仕,等.黄海南部小黄鱼生长、死亡和最适开捕体长[J].中国水产科学,2010,17(4):839-846.]
[66]Liu Z L,Xie H Y,Yan L P,et al.Comparative population dynamics of small yellow croaker Larimichthys polyactis in Southern Yellow Sea and East China Sea[J].Journal of Dalian Ocean University,2013,28(6):627-632.[刘尊雷,谢汉阳,严利平,等.黄海南部和东海小黄鱼资源动态的比较[J].大连海洋大学学报,2013,28(6):627-632.]
[67]Liu Q,Xu B D,Ye Z J,et.al.Growth and Mortality of small yellow croaker(Larimichthys polyactis)inhabiting Haizhou Bay of China[J].Journal of Ocean University of China,2012,11(4):557-561.
[68]Wang Y B,Yu C G,Yu C D,et al.Preliminary research on the rational utilization of Psenopsis anomala in the offshore water of South Zhejiang[J].Journal of Zhejiang Ocean University(Natural Science),2010,29(2):114-119.[王迎宾,俞存根,虞聪达,等.浙江南部外海刺鲳资源合理利用的初步研究[J].浙江海洋学院学报(自然科学版),2010,29(2):114-119.]
[69]Du J G,Lu Z B,Chen M R,et al.Changes in ecological parameters and resources of Japanese butterfish,Psenopsis anomala,in the middle and north Taiwan Strait[J].Journal of Oceanography in Taiwan Strait,2010,29(2):234-240.[杜建国,卢振彬,陈明茹,等.台湾海峡中北部海域刺鲳种群生态学参数及其变动趋势[J].台湾海峡,2010,29(2):234-240.]
[70]Zhang K,Chen Z Z,Qiu Y S.Decadal changes in growth,mortality and maturity parameters of Evynnis cardinalis in Beibu Gulf[J].South China Fisheries Science,2016,12(6):9-16.[张魁,陈作志,邱永松.北部湾二长棘犁齿鲷生长、死亡和性成熟参数的年际变化[J].南方水产科学,2016,12(6):9-16.].
[71]Hou G,Zhu X L,Lu H S.Growth,mortality and population composition of crimson sea bream,Paragyrops edita Tanaka in Beibu Gulf[J].Journal of Guangdong Ocean University,2008,28(3):50-55.[侯刚,朱立新,卢伙胜.北部湾二长棘鲷生长、死亡及其群体组成[J].广东海洋大学学报,2008,28(3):50-55.]
[72]Chen Z Z,Qiu Y S.Esitimation of growth and mortality parameters of Parargyrops edita Tanaka in Beibu Bay[J].Journal of Fisheries of China,2003,27(3):251-257.[陈作志,邱永松.北部湾二长棘鲷生长和死亡参数估计[J].水产学报,2003,27(3):251-257.]
[73]Feng B,Hou G,Lu H S.Management recommendation for Paragyrops edita in Beibu Gulf based on per-recruitment analysis[J].System Science and Comprehensive Studies in Agriculture,2009,25(1):114-119.[冯波,侯刚,卢伙胜.北部湾二长棘鲷资源开发保护策略研究[J].农业系统科学与综合研究,2009,25(1):114-119.]
[74]Wang X H,Qiu Y S,Du F Y.Study on the growth,mortality and optimum catchable size of Nemipterus virgatus in the Northern South China Sea[J].Journal of Ocean University of China,2004,34(2):224-230.[王雪辉,邱永松,杜飞雁.南海北部金线鱼生长、死亡和最适开捕体长研究[J].中国海洋大学学报,2004,34(2):224-230.]
[75]Wang X H,Qiu Y S,Du F Y.Study on the growth,mortality and optimum catchable size of Nemipterus Virgatus in the Northern South China Sea[J].Journal of Ocean University of China,2004,34(2):224-230.[王雪辉,邱永松,杜飞雁.南海北部金线鱼生长、死亡和最适开捕体长研究[J].中国海洋大学学报,2004,34(2):224-230.]
[76]Lu Z B,Du J G.Growth and mortality character of Nemi pterus virgatus in southern Taiwan Strait[J].Journal of Tropical Ocean Nography,2008,27(5):75-79.[卢振彬,杜建国.台湾海峡南部金线鱼的生长与死亡特性[J].热带海洋学报,2008,27(5):75-79.]
[77]Liu Y W,Xue Y,Wei B F,et al.An estimation of population parameters for five major fish species in the Haizhou Bay,Yellow Sea[J].Journal of Fishery Sciences of China,2014,21(1):125-133.[刘元文,薛莹,魏邦福,等.海州湾5种主要鱼类种群参数估算[J].中国水产科学,2014,21(1):125-133.]
[78]Liu J D,Lu H S,Zhu L X,et al.The difference of growth,mortality and group composition Between female and male Saurida tumbil in the Beibu Gulf[J].Marine Fisheries,2009,31(3):243-253.[刘金殿,卢伙胜,朱立新,等.北部湾多齿蛇鲻雌雄群体组成、生长、死亡特征的差异[J].海洋渔业,2009,31(3):243-253.]
[79]Shu L M,Qiu Y S.Biology analysis of Saurida tumbil in northern South China Sea[J].Journal of Fishery Sciences of China,2004,11(2):154-158.[舒黎明,邱永松,南海北部多齿蛇鲻生物学分析[J].中国水产科学,2004,11(2):154-158.]
[80]Liu Y,Cheng J H,Li S F.Utilization status of Setipinna taty in the East China Sea and its rational exploitation[J].Journal of Fishery Sciences of China,2006,13(3):485-491.[刘勇,程家骅,李圣法.东海区黄鲫资源的利用现状及合理利用探讨[J].中国水产科学,2006,13(3):485-491.]
[81]Lu Z B,Chen X.The parameters and their changes of the growth and mortality of Engraulidae and Clupeidae in Fujian coastal waters[J].Journal of Xiamen University(Natural Science),2008,47(2):279-285.[卢振彬,陈骁.福建沿海几种鲱、鳀科鱼类生长与死亡参数及其变化[J].厦门大学学报(自然科学版),2008,47(2):279-285.]
[82]Chen G B,Qiu Y S.Study on growth,mortality and reasonable utilization of Decapterus maruadsi in northern continental shelf waters of South China Sea[J].Journal of Oceanography in Taiwan Strait,2003,22(4):457-464.[陈国宝,邱永松.南海北部陆架区蓝圆鲹的生长、死亡及合理利用研究[J].台湾海峡,2003,22(4):457-464.]
[83]Zheng J,Wang Y B,Li R X,et al.Stock assessment of Jack Scomber japonicus and Decapterus maruadsi in coastal waters of Zhejiang Province[J].Journal of Zhejiang Ocean University(Natural Science),2012,31(4):309-315.[郑基,王迎宾,李仁星,等.浙江海域鲐鲹鱼资源量评估[J].浙江海洋学院学报(自然科学版),2012,31(4):309-315.]
[84]Costello C,Ovando D,Hilborn R,et al.Status and Solutions for the World’s Unassessed Fisheries[J].Science,2012,338(6106):517-520.
[85]Chen Y.Quality of fisheries data and uncertainty in stock assessment[J].Scientia Marina,2003,67(S1):75-87.