中国南海西沙群岛海域鸢乌贼角质颚生长特性
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摘要
角质颚是头足类重要的硬组织之一,被广泛用于研究类头足类渔业生物、生态学。根据我国灯光罩网渔船2016年1—3月和2017年1—3月在中国南海西沙群岛海域采集的818尾鸢乌贼样本,研究了鸢乌贼角质颚的外部形态生长特性。主成分分析表明,鸢乌贼的上头盖长(UHL)、上脊突长(UCL)、上翼长(UWL)、上侧壁长(ULWL)、下头盖长(LHL)、下脊突长(LCL)、下翼长(LWL)和下侧壁长(LLWL)可以作为鸢乌贼角质颚外形变化的特征参数。协方差分析表明:UHL、LWL与胴长的生长关系存在性别间显著性差异,其余参数均不存在显著性差异;所有外形特征参数与体质量间的生长关系均存在性别间显著性差异。AIC分析表明:除LLWL和雄性个体的UHL与胴长的关系最适合用线性方程表示外,其余均最适用指数方程表示;雌性个体所有外形参数与体质量的生长关系均最适合用对数函数表示,而雄性个体除UCL和UWL最适合用指数函数表示外,其余均最适合用线性方程表示。
The beak was one of the most important hard tissues of cephalopods which was widely used in the study of cephalopod fisheries biology and fishery ecology research. According to the measurement of the morphological parameters of the beak from 818 Sthenoteuthis oualaniensis samples collected by Chinese falling-net fishery during January to March of 2017 and 2018 in the Xisha Islands waters of the South China Sea, the morphological growth characteristic of the beak of S.oualaniensis was studied. The principal component analysis showed that the upper hood length(UHL), the upper crest length(UCL), the upper wing length(UWL), the upper lateral wall length(ULWL), the lower hood length(LHL), the lower crest length(LCL),the lower wing length(LWL) and the lower lateral wall length(LLWL) could be used as themorphological characteristic parameters to study the morphological changes of the beak. The analysis of covariance(ANCOVA) showed that there were significant differences in the relationships between UHL, LWL versus mantel length(ML) between males and females, but no significant difference was found in other characteristic parameters. The relationships between beak morphological variables and body weight(BW) differed significantly between males and females in all of the morphological characteristic parameters. The relationships between LLWL,UHLof males and ML were optimally described by the linear growth models, and exponential functions were optimal for the other parameters. The logarithmic functions were optimal for the relationship between the ULWL, LHL, LCL, LWL, LLWL and BW for females, however exponential equations were optimal for the UCL,UWL, and linear functions were optimal for the other parameters for males.
The beak was one of the most important hard tissues of cephalopods which was widely used in the study of cephalopod fisheries biology and fishery ecology research. According to the measurement of the morphological parameters of the beak from 818 Sthenoteuthis oualaniensis samples collected by Chinese falling-net fishery during January to March of 2017 and 2018 in the Xisha Islands waters of the South China Sea, the morphological growth characteristic of the beak of S.oualaniensis was studied. The principal component analysis showed that the upper hood length(UHL), the upper crest length(UCL), the upper wing length(UWL), the upper lateral wall length(ULWL), the lower hood length(LHL), the lower crest length(LCL),the lower wing length(LWL) and the lower lateral wall length(LLWL) could be used as themorphological characteristic parameters to study the morphological changes of the beak. The analysis of covariance(ANCOVA) showed that there were significant differences in the relationships between UHL, LWL versus mantel length(ML) between males and females, but no significant difference was found in other characteristic parameters. The relationships between beak morphological variables and body weight(BW) differed significantly between males and females in all of the morphological characteristic parameters. The relationships between LLWL,UHLof males and ML were optimally described by the linear growth models, and exponential functions were optimal for the other parameters. The logarithmic functions were optimal for the relationship between the ULWL, LHL, LCL, LWL, LLWL and BW for females, however exponential equations were optimal for the UCL,UWL, and linear functions were optimal for the other parameters for males.
引文
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[7] 招春旭,陈昭澎,何雄波,等.基于耳石微结构的南海春季鸢乌贼日龄、生长与种群结构的研究[J].水生生物学报,2017,41(4):884-890.ZHAO C X,CHEN Z P,HE X B,et al.Age,growth and population structure of purple back flying squid,Sthenoteuthis oulaniensis in the South China Sea in spring based on statolith microstructure[J].Acta Hydrobiologica Sinica,2017,41(4):884-890.
[8] 陆化杰,王从军,陈新军.4—6月东太平洋赤道公海鸢乌贼生物学特性初步研究[J].上海海洋大学学报,2014,23(3):441-447.LU H J,WANG C J,CHEN X J.Preliminary study on the biological characteristics of Sthenoteuthis oualaniensis in the high seas nearby the equator of eastern Pacific during April to June[J].Journal of Shanghai Ocean University,2014,23(3):441-447.
[9] CLARKE M R.The identification of cephalopod Z‘beaks’ and the relationship between break size and total body weight[J].Bulletin of the British Museum of Natural History,Zoology,1962,8(10):419-480.
[10] HERNáNDEZ-LóPEZ J L,CASTRO-HERNANDEZ J L,HERNáNDEZ-GARCIA V.Age determined from the daily deposition of concentric rings on common octopus (Octopus vulgaris) beaks[J].Fishery Bulletin,2001,99(4):679-684.
[11] 陈芃,方舟,陈新军.基于角质颚外部形态学的柔鱼种群判别[J].海洋渔业,2015,37(1):1-9.CHEN P,FANG Z,CHEN X J.Population identification of Ommastrephes bartramii based on beak’s external morphological characters[J].Marine Fisheries,2015,37(1):1-9.
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[14] 许嘉锦.Octopus与Cistpous属章鱼口器地标点之几何形态学研究[D].台北:中山大学海洋生物研究所,2003.HSU C C.Geomorphometric study of Octopus and Cistopus (Cephalopoda:Octopodidae) based on landmarks of beaks[D].Taipei:Institute of Marine Biology,Central University,2003.
[15] 胡贯宇,金岳,陈新军.秘鲁外海茎柔鱼角质颚的形态特征及其与个体大小和日龄的关系[J].海洋渔业,2017,39(4):361-371.HU G Y,JIN Y,CHEN X J.Beak morphological characteristics of Dosidicus gigas off the Peruvian Exclusive Economic Zone(EEZ) and their relationship with body size and daily age[J].Marine Fisheries,2017,39(4):361-371.
[16] JACKSON G D,MCKINNON J F.Beak length analysis of arrow squid Nototodarus sloanii (Cephalopoda:Ommastrephidae) in southern New Zealand waters[J].Polar Biology,1996,16(3):227-230.
[17] MALCOLM H.Modelling and quantitative methods in fisheries[M].2nd ed.Florida:Chapman & Hall/CRC,2001:227-232.
[18] HIRAMATSU K.Application of maximum likelihood method and AIC to fish population dynamics[M]//MATSUMIYA Y.Fish Population Dynamics and Statistical Models.Tokyo:Koseisha Koseikaku,1993:9-21.
[19] IMAI C,SAKAI H,KATSURA K,et al.Growth model for the endangered cyprinid fish Tribolodon nakamurai based on otolith analyses[J].Fisheries Science,2002,68(4):843-848.
[20] 金岳,方舟,李云凯,等.北太平洋东部柔鱼群体角质颚生长特性分析[J].海洋渔业,2015,37(2):101-106.JIN Y,FANG Z,LI Y K,et al.Growth characteristics analysis of Ommastrephes bartramii beaks in eastern North Pacific[J].Marine Fisheries,2015,37(2):101-106.
[21] 方舟,陈新军,陆化杰,等.北太平洋两个柔鱼群体角质颚形态及生长特征[J].生态学报,2014,34(19):5405-5415.FANG Z,CHEN X J,LU H J,et al.Morphology and growth of beaks in two cohorts for neon flying squid (Ommastrephes bartramii) in the North Pacific Ocean[J].Acta Ecologica Sinica,2014,34(19):5405-5415.
[22] CHEN X J,LU H J,LIU B L,et al.Species identifcation of Ommastrephes bartramii,Dosidicus gigas,Sthenoteuthis oualaniensis and Illex argentinus (Ommastrephidae) by using beak morphological variables[J].Scientia Marina,2012,76(3):473-481.
[23] 刘必林,陈新军.印度洋西北海域鸢乌贼角质颚长度分析[J].渔业科学进展,2010,31(1):8-14.LIU B L,CHEN X J.Beak length analysis of the purpleback flying squid Sthenoeuthis oualaniensis in northwest Indian Ocean[J].Progress in Fishery Sciences,2010,31(1):8-14.
[24] 徐杰,刘尊雷,李圣法,等.东海剑尖枪乌贼角质颚的外部形态及生长特性[J].海洋渔业,2016,38(3):245-253.XU J,LIU Z L,LI S F,et al.Morphology and growth characteristics of Uroteuthis edulis beak in the East China Sea[J].Marine Fisheries,2016,38(3):245-253.
[25] 杨林林,姜亚洲,刘尊雷,等.东海火枪乌贼角质颚的形态特征[J].中国水产科学,2012,19(4):586-593.YANG L L,JIANG Y Z,LIU Z L,et al.Analysis of beak morphology of Loligo beka in the East China Sea[J].Journal of Fishery Sciences of China,2012,19(4):586-593.
[26] 杨林林,姜亚洲,刘尊雷,等.东海太平洋褶柔鱼角质颚的形态学分析[J].中国海洋大学学报,2012,42(10):51-57.YANG L L,JIANG Y Z,LIU Z L,et al.Variation analysis on partial morphometric measurements of beak of Todarodes pacificus inhabiting East China Sea[J].Journal of Ocean University of China,2012,42(10):51-57.
[27] QUEIRóS J P,CHEREL Y,CEIA F R,et al.Ontogenic changes in habitat and trophic ecology in the Antarctic squid Kondakovia longimana derived from isotopic analysis on beaks[J].Polar Biology,2018,41(12):2409-2421.
[28] GUERREIRO M,PHILLIPS R A,CHEREL Y,et al.Habitat and trophic ecology of Southern Ocean cephalopods from stable isotope analyses[J].Marine Ecology Progress Series,2015,530:119-134.
[29] ARKHIPKIN A,LAPTIKHOVSKY V.Seasonal and interannual variability in growth and maturation of winter-spawning Illex argentinus (Cephalopoda,Ommastrephidae) in the Southwest Atlantic[J].Aquatic Living Resources,1994,7(4):221-232.
[30] CHEN X J,LU H J,LIU B L,et al.Relationship between beak morphological variables and body size and mantle length of male and female Argentine shortfin squid (Illex argentinus)[J].Journal of Ocean University of China,2012,11(4):539-546.
[2] 陈新军,韩宝平,刘必林,等.世界头足类资源及其渔业[M].北京:科学出版社,2013:28-31.CHEN X J,HAN B P,LIU B L,et al.The resources and fisheries of cephalopod in the world[M].Beijing:Science Press,2013:28-31.
[3] 张鹏,杨吝,张旭丰,等.南海金枪鱼和鸢乌贼资源开发现状及前景[J].南方水产,2010,6(1):68-74.ZHANG P,YANG L,ZHANG X F,et al.The present status and prospect on exploitotion of tuna and squid fishery resources in South China Sea[J].South China Fisheries Science,2010,6(1):68-73.
[4] 贾晓平,李永振,李纯厚,等.南海专属经济区和大陆架渔业生态环境与渔业资源[M].北京:科学出版社,2004:385-391.JIA X P,LI Y Z,LI C H,et al.Fishery ecological environment and fishery resources in the exclusive economic zone and continental shelf of the South China Sea[M].Beijing:Science Press,2004:385-391.
[5] CHEN X J,LIU B L,TIAN S Q,et al.Fishery biology of purpleback squid,Sthenoteuthis oualaniensis,in the northwest Indian Ocean[J].Fisheries Research,2007,83(1):98-104.
[6] JIN Y,LIU B L,CHEN X J,et al.Morphological beak differences of loliginid squid,Uroteuthis chinensis and Uroteuthis edulis,in the northern South China Sea[J].Journal of Oceanology and Limnology,2018,36(2):559-571.
[7] 招春旭,陈昭澎,何雄波,等.基于耳石微结构的南海春季鸢乌贼日龄、生长与种群结构的研究[J].水生生物学报,2017,41(4):884-890.ZHAO C X,CHEN Z P,HE X B,et al.Age,growth and population structure of purple back flying squid,Sthenoteuthis oulaniensis in the South China Sea in spring based on statolith microstructure[J].Acta Hydrobiologica Sinica,2017,41(4):884-890.
[8] 陆化杰,王从军,陈新军.4—6月东太平洋赤道公海鸢乌贼生物学特性初步研究[J].上海海洋大学学报,2014,23(3):441-447.LU H J,WANG C J,CHEN X J.Preliminary study on the biological characteristics of Sthenoteuthis oualaniensis in the high seas nearby the equator of eastern Pacific during April to June[J].Journal of Shanghai Ocean University,2014,23(3):441-447.
[9] CLARKE M R.The identification of cephalopod Z‘beaks’ and the relationship between break size and total body weight[J].Bulletin of the British Museum of Natural History,Zoology,1962,8(10):419-480.
[10] HERNáNDEZ-LóPEZ J L,CASTRO-HERNANDEZ J L,HERNáNDEZ-GARCIA V.Age determined from the daily deposition of concentric rings on common octopus (Octopus vulgaris) beaks[J].Fishery Bulletin,2001,99(4):679-684.
[11] 陈芃,方舟,陈新军.基于角质颚外部形态学的柔鱼种群判别[J].海洋渔业,2015,37(1):1-9.CHEN P,FANG Z,CHEN X J.Population identification of Ommastrephes bartramii based on beak’s external morphological characters[J].Marine Fisheries,2015,37(1):1-9.
[12] LU C C,ICKERINGILL R.Cephalopod beak identification and biomass estimation techniques:tools for dietary studies of southern Australian finfishes[R].Victoria:Museum Victoria Science,2002.
[13] 陆化杰,陈新军,方舟.西南大西洋阿根廷滑柔鱼2个不同产卵群间角质颚外形生长特性比较[J].中国海洋大学学报,2012,42(10):33-40.LU H J,CHEN X J,FANG Z.Comparison of the beak morphologic growth characteristics between two spawning populations of Illex argentinus in southwest Atlantic Ocean[J].Journal of Ocean University of China,2012,42(10):33-40.
[14] 许嘉锦.Octopus与Cistpous属章鱼口器地标点之几何形态学研究[D].台北:中山大学海洋生物研究所,2003.HSU C C.Geomorphometric study of Octopus and Cistopus (Cephalopoda:Octopodidae) based on landmarks of beaks[D].Taipei:Institute of Marine Biology,Central University,2003.
[15] 胡贯宇,金岳,陈新军.秘鲁外海茎柔鱼角质颚的形态特征及其与个体大小和日龄的关系[J].海洋渔业,2017,39(4):361-371.HU G Y,JIN Y,CHEN X J.Beak morphological characteristics of Dosidicus gigas off the Peruvian Exclusive Economic Zone(EEZ) and their relationship with body size and daily age[J].Marine Fisheries,2017,39(4):361-371.
[16] JACKSON G D,MCKINNON J F.Beak length analysis of arrow squid Nototodarus sloanii (Cephalopoda:Ommastrephidae) in southern New Zealand waters[J].Polar Biology,1996,16(3):227-230.
[17] MALCOLM H.Modelling and quantitative methods in fisheries[M].2nd ed.Florida:Chapman & Hall/CRC,2001:227-232.
[18] HIRAMATSU K.Application of maximum likelihood method and AIC to fish population dynamics[M]//MATSUMIYA Y.Fish Population Dynamics and Statistical Models.Tokyo:Koseisha Koseikaku,1993:9-21.
[19] IMAI C,SAKAI H,KATSURA K,et al.Growth model for the endangered cyprinid fish Tribolodon nakamurai based on otolith analyses[J].Fisheries Science,2002,68(4):843-848.
[20] 金岳,方舟,李云凯,等.北太平洋东部柔鱼群体角质颚生长特性分析[J].海洋渔业,2015,37(2):101-106.JIN Y,FANG Z,LI Y K,et al.Growth characteristics analysis of Ommastrephes bartramii beaks in eastern North Pacific[J].Marine Fisheries,2015,37(2):101-106.
[21] 方舟,陈新军,陆化杰,等.北太平洋两个柔鱼群体角质颚形态及生长特征[J].生态学报,2014,34(19):5405-5415.FANG Z,CHEN X J,LU H J,et al.Morphology and growth of beaks in two cohorts for neon flying squid (Ommastrephes bartramii) in the North Pacific Ocean[J].Acta Ecologica Sinica,2014,34(19):5405-5415.
[22] CHEN X J,LU H J,LIU B L,et al.Species identifcation of Ommastrephes bartramii,Dosidicus gigas,Sthenoteuthis oualaniensis and Illex argentinus (Ommastrephidae) by using beak morphological variables[J].Scientia Marina,2012,76(3):473-481.
[23] 刘必林,陈新军.印度洋西北海域鸢乌贼角质颚长度分析[J].渔业科学进展,2010,31(1):8-14.LIU B L,CHEN X J.Beak length analysis of the purpleback flying squid Sthenoeuthis oualaniensis in northwest Indian Ocean[J].Progress in Fishery Sciences,2010,31(1):8-14.
[24] 徐杰,刘尊雷,李圣法,等.东海剑尖枪乌贼角质颚的外部形态及生长特性[J].海洋渔业,2016,38(3):245-253.XU J,LIU Z L,LI S F,et al.Morphology and growth characteristics of Uroteuthis edulis beak in the East China Sea[J].Marine Fisheries,2016,38(3):245-253.
[25] 杨林林,姜亚洲,刘尊雷,等.东海火枪乌贼角质颚的形态特征[J].中国水产科学,2012,19(4):586-593.YANG L L,JIANG Y Z,LIU Z L,et al.Analysis of beak morphology of Loligo beka in the East China Sea[J].Journal of Fishery Sciences of China,2012,19(4):586-593.
[26] 杨林林,姜亚洲,刘尊雷,等.东海太平洋褶柔鱼角质颚的形态学分析[J].中国海洋大学学报,2012,42(10):51-57.YANG L L,JIANG Y Z,LIU Z L,et al.Variation analysis on partial morphometric measurements of beak of Todarodes pacificus inhabiting East China Sea[J].Journal of Ocean University of China,2012,42(10):51-57.
[27] QUEIRóS J P,CHEREL Y,CEIA F R,et al.Ontogenic changes in habitat and trophic ecology in the Antarctic squid Kondakovia longimana derived from isotopic analysis on beaks[J].Polar Biology,2018,41(12):2409-2421.
[28] GUERREIRO M,PHILLIPS R A,CHEREL Y,et al.Habitat and trophic ecology of Southern Ocean cephalopods from stable isotope analyses[J].Marine Ecology Progress Series,2015,530:119-134.
[29] ARKHIPKIN A,LAPTIKHOVSKY V.Seasonal and interannual variability in growth and maturation of winter-spawning Illex argentinus (Cephalopoda,Ommastrephidae) in the Southwest Atlantic[J].Aquatic Living Resources,1994,7(4):221-232.
[30] CHEN X J,LU H J,LIU B L,et al.Relationship between beak morphological variables and body size and mantle length of male and female Argentine shortfin squid (Illex argentinus)[J].Journal of Ocean University of China,2012,11(4):539-546.