我国几种常见头足类角质颚碳氮稳定同位素分析
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摘要
头足类角质颚具有稳定的化学成分和物理结构,其稳定同位素分析已成为头足类生态学研究的主要方法之一。本文分别对4种近海头足类(曼氏无针乌贼、杜氏枪乌贼、短蛸和剑尖枪乌贼)角质颚进行稳定同位素分析,研究角质颚中碳、氮稳定同位素比值(~(13)C/~(12)C,δ~(13)C;~(15)N/~(14)N,δ~(15)N)、摄食生态和栖息环境在物种间的差异,分析可能形成差异的原因以及δ~(13)C、δ~(15)N值与头足类胴长间的相关性,旨在解读不同头足类所生活栖息的渔场,了解它们所处生态系统中的地位,为渔业开发与管理提供材料,同时也为我们可以更好地利用这些物种提供基础。结果表明,4种头足类的平均胴长关系为杜氏枪乌贼>剑尖枪乌贼>曼氏无针乌贼>短蛸(122.72 mm>67.93 mm>67.93 mm>35.3 mm),平均体质量关系为杜氏枪乌贼>曼氏无针乌贼>短蛸>剑尖枪乌贼(70.39 g>50.67 g>47.35 g>19.93 g),其中:曼氏无针乌贼的δ~(15)N值最大(10.81‰±0.38‰),δ~(13)C值最小(-14.98‰±0.55‰),C/N比值最小(3.08±0.03);剑尖枪乌贼的δ~(15)N值最小(8.21‰±0.30‰),δ~(13)C值最大(-13.00‰±0.54‰),C/N比值最大(3.57±0.06)。曼氏无针乌贼可能具有较高营养水平,剑尖枪乌贼、杜氏枪乌贼相较于短蛸、曼氏无针乌贼生活在较浅水域的栖息环境。杜氏枪乌贼与短蛸的δ~(13)C、δ~(15)N值无显著差异(ANOVA,P>0.05),它们的生态位圆存在重叠,处于相近的营养级,其他物种间角质颚的δ~(13)C、δ~(15)N值存在显著差异(ANOVA,P<0.05),生态位圆无重叠。此外杜氏枪乌贼角质颚的δ~(13)C、δ~(15)N值与胴长(ML)均呈显著的线性关系(δ~(13)C=0.023 6×L_(ML)-17.429,P<0.05;δ~(15)N=0.017 6×L_(ML)+6.873,P<0.05),而曼氏无针乌贼、短蛸以及剑尖枪乌贼的胴长与角质颚的δ~(13)C、δ~(15)N值相关性不显著(P>0.05),说明其摄食习性随体长变化无显著相关。
The beaks of cephalopods have stable chemical compositions and firm physical structures. Recently, with the increasing use of stable isotope analysis in ecology, it becomes a powerful tool and complement of traditional methods for investigating the trophic ecology and migration patterns of invertebrates. This research was based on 18 Uroteuthis duvauceli specimens, 18 Sepiella maindronis specimens, 20 Octopus ocellatus specimens and 15 Uroteuthis edulis specimens, which were collected in August 2013 from the East China Sea. The beaks of these four kinds of offshore species were studied by determining their carbon stable isotope(~(13)C/~(12)C,δ~(13)C)and nitrogen stable isotope(~(15)N/~(14)N,δ~(15)N).In order to find the differences of these species' habitats and the foraging behaviors, the δ~(13)C and δ~(15)N values were analyzed to see whether there was a correlation with their mantle lengths. The study aimed to explain the habitats of different cephalopods and find their position in the ecological system, not only providing materials for further development and management, but also giving a basis for the better use of these species. The results showed that the average values of mantle length for four species: U.duvauceli>U.edulis>S.maindroni>O.ocellatus(122.72 mm>67.93 mm >67.93 mm >35.3 mm). The average values of body weight for four species: U.duvauceli>S.maindroni>O.ocellatus>U.edulis>(70.39 g>50.67 g >47.35 g >19.93 g). S.maindroni have the maximum δ~(15)N value(10.81‰±0.38‰),the minimum δ~(13)C value(-14.98‰±0.55‰),the minimum C/N(3.08±0.03)and U.edulishave the minimum δ~(15)N value(8.21‰±0.30‰), the maximum δ~(13)C value(-13.00‰±0.54‰), the maximum C/N(3.57±0.06), U.duvauceli and U.edulislive in the shallower water area than the S.maindroni and O.ocellatus. And the δ~(13)C and δ~(15)N values between U.duvauceli and O.ocellatus were very similar(ANOVA, P> 0.05), their ecological niches were much overlapped, they may have the similar trophic level. The δ~(13)C and δ~(15)N values of other kinds of species had significant differences(ANOVA, P< 0.05). In addition, the δ~(13)C values of U.duvauceli and the mantle length(ML) showed a significant liner relationship(δ~(13)C=0.023 6×L_(ML)-17.429,P< 0.05), δ~(15)N values also showed a significant liner relationship with the mantle length(ML)(δ~(15)N =0.017 6×L_(ML)+ 6.873,P<0.05), but there was no significant correlation between carbon and nitrogen stable isotopes with the mantle length in S.maindroni, O.ocellatus and U.edulis(P > 0.05).It showed that there was no significant connection between feeding habit and mantle length.
The beaks of cephalopods have stable chemical compositions and firm physical structures. Recently, with the increasing use of stable isotope analysis in ecology, it becomes a powerful tool and complement of traditional methods for investigating the trophic ecology and migration patterns of invertebrates. This research was based on 18 Uroteuthis duvauceli specimens, 18 Sepiella maindronis specimens, 20 Octopus ocellatus specimens and 15 Uroteuthis edulis specimens, which were collected in August 2013 from the East China Sea. The beaks of these four kinds of offshore species were studied by determining their carbon stable isotope(~(13)C/~(12)C,δ~(13)C)and nitrogen stable isotope(~(15)N/~(14)N,δ~(15)N).In order to find the differences of these species' habitats and the foraging behaviors, the δ~(13)C and δ~(15)N values were analyzed to see whether there was a correlation with their mantle lengths. The study aimed to explain the habitats of different cephalopods and find their position in the ecological system, not only providing materials for further development and management, but also giving a basis for the better use of these species. The results showed that the average values of mantle length for four species: U.duvauceli>U.edulis>S.maindroni>O.ocellatus(122.72 mm>67.93 mm >67.93 mm >35.3 mm). The average values of body weight for four species: U.duvauceli>S.maindroni>O.ocellatus>U.edulis>(70.39 g>50.67 g >47.35 g >19.93 g). S.maindroni have the maximum δ~(15)N value(10.81‰±0.38‰),the minimum δ~(13)C value(-14.98‰±0.55‰),the minimum C/N(3.08±0.03)and U.edulishave the minimum δ~(15)N value(8.21‰±0.30‰), the maximum δ~(13)C value(-13.00‰±0.54‰), the maximum C/N(3.57±0.06), U.duvauceli and U.edulislive in the shallower water area than the S.maindroni and O.ocellatus. And the δ~(13)C and δ~(15)N values between U.duvauceli and O.ocellatus were very similar(ANOVA, P> 0.05), their ecological niches were much overlapped, they may have the similar trophic level. The δ~(13)C and δ~(15)N values of other kinds of species had significant differences(ANOVA, P< 0.05). In addition, the δ~(13)C values of U.duvauceli and the mantle length(ML) showed a significant liner relationship(δ~(13)C=0.023 6×L_(ML)-17.429,P< 0.05), δ~(15)N values also showed a significant liner relationship with the mantle length(ML)(δ~(15)N =0.017 6×L_(ML)+ 6.873,P<0.05), but there was no significant correlation between carbon and nitrogen stable isotopes with the mantle length in S.maindroni, O.ocellatus and U.edulis(P > 0.05).It showed that there was no significant connection between feeding habit and mantle length.
引文
[1] 刘必林,陈新军.头足类角质颚的研究进展[J].水产学报,2009,33(1):157-164.LIU B L,CHEN X J.Review on the research development of beaks in Cephalopoda[J].Journal of Fisheries of China,2009,33(1):157-164.
[2] CLARKE M R.The identification of cephalopod “beaks” and the relationship between beak size and total body weight[J].Bulletin of the British Museum (Natural History).Zoology,1962,8(10):419-480.
[3] CLARKE M R.A handbook for the identification of cephalopod beaks[M].Oxford:Clarendon Press,1986:273.
[4] CHEN X J,LU H J,LIU B L,et al.Species identification of Ommastrephes bartramii,Dosidicus gigas,Sthenoteuthis oualaniensis and Illex argentinus (Ommastrephidae) using beak morphological variables[J].Scientia Marina,2012,76(3):473-481.
[5] LIU B L,FANG Z,CHEN X J,et al.Spatial variations in beak structure to identify potentially geographic populations of Dosidicus gigas in the eastern Pacific Ocean[J].Fisheries Research,2015,164:185-192.
[6] CLARKE M R,TRILLMICH F.Cephalopods in the diet of fur seals of the Galapagos Islands[J].Journal of Zoology,1980,190(2):211-215.
[7] SEKIGUCHI K,KLAGES N T W,BEST P B.The diet of strap-toothed whales (Mesoplodon layardii)[J].Journal of Zoology,1996,239(3):453-463.
[8] HERNáNDEZ-LóPEZ J L,CASTRO-HERNáNDEZ J L.Age determined from the daily deposition of concentric rings on common octopus (Octopus vulgaris) beaks[J].Fishery Bulletin,2001,99(4):679-684.
[9] JACKSON G D.The use of beaks as tools for biomass estimation in the deepwater squid Moroteuthis ingens (Cephalopoda:Onychoteuthidae) in New Zealand waters[J].Polar Biology,1995,15(1):9-14.
[10] AUDZIJONYTE A,FULTON E A,KUPARINEN A.Corrections:Stable carbon isotopes and carbon flow in ecosystems[J].BioScience,1986,36(7):475.
[11] POST D M.Using stable isotopes to estimate trophic position:models,methods,and assumptions[J].Ecology,2002,83(3):703-718.
[12] 隋昊志,韩东燕,薛莹,等.基于碳、氮稳定同位素研究胶州湾普氏栉虾虎鱼的摄食习性[J].应用生态学报,2017,28(11):3789-3796.SUI H Z,HAN D Y,XUE Y,et al.Feeding habits of Rhinogobius pflaumi in Jiaozhou Bay,China based on carbon and nitrogen stable isotope analysis[J].Chinese Journal of Applied Ecology,2017,28(11):3789-3796.
[13] 蔡德陵,李红燕,唐启升,等.黄东海生态系统食物网连续营养谱的建立:来自碳氮稳定同位素方法的结果[J].中国科学 C辑生命科学,2005,35(2):123-130.CAI D L,LI H Y,TANG Q S,et al.Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem:Insight from carbon and nitrogen stable isotopes[J].Science in China Series C Life Sciences,2005,48(6):531-539.
[14] 李忠义,金显仕,庄志猛,等.稳定同位素技术在水域生态系统研究中的应用[J].生态学报,2005,25(11):3052-3060.LI Z Y,JIN X S,ZHUANG Z M,et al.Applications of stable isotope techniques in aquatic ecological studies[J].Acta Ecologica Sinica,2005,25(11):3052-3060.
[15] OVERMAN N C,PARRISH D L.Stable isotope composition of walleye:15N accumulation with age and area-specific differences in δ13C[J].Canadian Journal of Fisheries and Aquatic Sciences,2001,58(6):1253-1260.
[16] SMITH K A.Soil analysis:modern instrumental techniques[M].2nd ed.New York:Marcel Dekker,1991.
[17] ADAMS D C,OTáROLA-CASTILLO E.Geomorph:an R package for the collection and analysis of geometric morphometric shape data[J].Methods in Ecology and Evolution,2013,4(4):393-399.
[18] LAYMAN C A,ARRINGTON D A,MONTANA C G,et al.Can stable isotope ratios provide for community-wide measures of trophic structure[J].Ecology,2007,88(1):42-48.
[19] HOBSON K A,PIATT J F,PITOCCHELLI J.Using stable isotopes to determine seabird trophic relationships[J].Journal of Animal Ecology,1994,63(4):786-798.
[20] CHEREL Y,HOBSON K A.Geographical variation in carbon stable isotope signatures of marine predators:a tool to investigate their foraging areas in the southern Ocean[J].Marine Ecology Progress Series,2007,329:281-287.
[21] TAKAI N,ONAKA S,IKEDA Y,et al.Geographical variations in carbon and nitrogen stable isotope ratios in squid[J].Journal of the Marine Biological Association of the United Kingdom,2000,80(4):675-684.
[22] 陈新军,刘必林,王尧耕.世界头足类[M].北京:海洋出版社,2009.CHEN X J,LIU B L,WANG Y G.Cephalopods of the world[M].Beijing:Ocean Press,2009.
[23] DENIRO M J,EPSTEIN S.Influence of diet on the distribution of carbon isotopes in animals[J].Geochimica et Cosmochimica Acta,1978,42(5):495-506.
[24] RUIZ-COOLEY R I,MARKAIDA U,GENDRON D,et al.Stable isotopes in jumbo squid (Dosidicus gigas) beaks to estimate its trophic position:comparison between stomach contents and stable isotopes[J].Journal of the Marine Biological Association of the United Kingdom,2006,86(2):437-445.
[25] NEGRI A,DANERI G A,CEIA F,et al.The cephalopod prey of the Weddell seal,Leptonychotes weddellii,a biological sampler of the Antarctic marine ecosystem[J].Polar Biology,2016,39(3):561-564.
[26] 方舟,金岳,胡飞飞,等.秋季我国近海不同海域短蛸角质颚形态学研究[J].水产学报,2018,42(7):1050-1059.FANG Z,JIN Y,HU F F,et al.Beak morphometrics of short arm octopus (Amphioctopus fangsiao) in different sea areas of China in autumn[J].Journal of Fisheries of China,2018,42(7):1050-1059.
[27] GRAHAM B S,KOCH P L,NEWSOME S D,et al.Using Isoscapes to trace the movements and foraging behavior of top predators in oceanic ecosystems[M]//WEST J B,BOWEN G J,DAWSON T E,et al.Isoscapes:Understanding Movement,Pattern,and Process on Earth through Isotope Mapping.New York:Springer,2010:299-318.
[28] 金岳,陈新军,李云凯,等.基于稳定同位素技术的北太平洋柔鱼角质颚信息[J].生态学杂志,2014,33(8):2101-2107.JIN Y,CHEN X J,LI Y K,et al.Information in beaks of Ommastrephes bartramii in the North Pacific Ocean using stable isotope technology[J].Chinese Journal of Ecology,2014,33(8):2101-2107.
[29] RAU G H,SWEENEY R E,KAPLAN I R.Plankton 13C∶12C ratio changes with latitude:differences between northern and southern oceans[J].Deep Sea Research Part A:Oceanographic Research Papers,1982,29(8):1035-1039.
[30] CHEREL Y,RIDOUX V,SPITZ J,et al.Stable isotopes document the trophic structure of a deep-sea cephalopod assemblage including giant octopod and giant squid[J].Biology Letters,2009,5(3):364-367.
[31] FANG Z,THOMPSON K,JIN Y,et al.Preliminary analysis of beak stable isotopes (δ13C and δ15N) stock variation of neon flying squid,Ommastrephes bartramii,in the North Pacific Ocean[J].Fisheries Research,2016,177:153-163.
[32] LU C C,ICKERINGILL R.Cephalopod beak identification and biomass estimation techniques:tools for dietary studies of southern Australian finfishes[R].Museum Victoria Science Reports.Australia:Fisheries Research and Development Corporation,2002:1-65.
[2] CLARKE M R.The identification of cephalopod “beaks” and the relationship between beak size and total body weight[J].Bulletin of the British Museum (Natural History).Zoology,1962,8(10):419-480.
[3] CLARKE M R.A handbook for the identification of cephalopod beaks[M].Oxford:Clarendon Press,1986:273.
[4] CHEN X J,LU H J,LIU B L,et al.Species identification of Ommastrephes bartramii,Dosidicus gigas,Sthenoteuthis oualaniensis and Illex argentinus (Ommastrephidae) using beak morphological variables[J].Scientia Marina,2012,76(3):473-481.
[5] LIU B L,FANG Z,CHEN X J,et al.Spatial variations in beak structure to identify potentially geographic populations of Dosidicus gigas in the eastern Pacific Ocean[J].Fisheries Research,2015,164:185-192.
[6] CLARKE M R,TRILLMICH F.Cephalopods in the diet of fur seals of the Galapagos Islands[J].Journal of Zoology,1980,190(2):211-215.
[7] SEKIGUCHI K,KLAGES N T W,BEST P B.The diet of strap-toothed whales (Mesoplodon layardii)[J].Journal of Zoology,1996,239(3):453-463.
[8] HERNáNDEZ-LóPEZ J L,CASTRO-HERNáNDEZ J L.Age determined from the daily deposition of concentric rings on common octopus (Octopus vulgaris) beaks[J].Fishery Bulletin,2001,99(4):679-684.
[9] JACKSON G D.The use of beaks as tools for biomass estimation in the deepwater squid Moroteuthis ingens (Cephalopoda:Onychoteuthidae) in New Zealand waters[J].Polar Biology,1995,15(1):9-14.
[10] AUDZIJONYTE A,FULTON E A,KUPARINEN A.Corrections:Stable carbon isotopes and carbon flow in ecosystems[J].BioScience,1986,36(7):475.
[11] POST D M.Using stable isotopes to estimate trophic position:models,methods,and assumptions[J].Ecology,2002,83(3):703-718.
[12] 隋昊志,韩东燕,薛莹,等.基于碳、氮稳定同位素研究胶州湾普氏栉虾虎鱼的摄食习性[J].应用生态学报,2017,28(11):3789-3796.SUI H Z,HAN D Y,XUE Y,et al.Feeding habits of Rhinogobius pflaumi in Jiaozhou Bay,China based on carbon and nitrogen stable isotope analysis[J].Chinese Journal of Applied Ecology,2017,28(11):3789-3796.
[13] 蔡德陵,李红燕,唐启升,等.黄东海生态系统食物网连续营养谱的建立:来自碳氮稳定同位素方法的结果[J].中国科学 C辑生命科学,2005,35(2):123-130.CAI D L,LI H Y,TANG Q S,et al.Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem:Insight from carbon and nitrogen stable isotopes[J].Science in China Series C Life Sciences,2005,48(6):531-539.
[14] 李忠义,金显仕,庄志猛,等.稳定同位素技术在水域生态系统研究中的应用[J].生态学报,2005,25(11):3052-3060.LI Z Y,JIN X S,ZHUANG Z M,et al.Applications of stable isotope techniques in aquatic ecological studies[J].Acta Ecologica Sinica,2005,25(11):3052-3060.
[15] OVERMAN N C,PARRISH D L.Stable isotope composition of walleye:15N accumulation with age and area-specific differences in δ13C[J].Canadian Journal of Fisheries and Aquatic Sciences,2001,58(6):1253-1260.
[16] SMITH K A.Soil analysis:modern instrumental techniques[M].2nd ed.New York:Marcel Dekker,1991.
[17] ADAMS D C,OTáROLA-CASTILLO E.Geomorph:an R package for the collection and analysis of geometric morphometric shape data[J].Methods in Ecology and Evolution,2013,4(4):393-399.
[18] LAYMAN C A,ARRINGTON D A,MONTANA C G,et al.Can stable isotope ratios provide for community-wide measures of trophic structure[J].Ecology,2007,88(1):42-48.
[19] HOBSON K A,PIATT J F,PITOCCHELLI J.Using stable isotopes to determine seabird trophic relationships[J].Journal of Animal Ecology,1994,63(4):786-798.
[20] CHEREL Y,HOBSON K A.Geographical variation in carbon stable isotope signatures of marine predators:a tool to investigate their foraging areas in the southern Ocean[J].Marine Ecology Progress Series,2007,329:281-287.
[21] TAKAI N,ONAKA S,IKEDA Y,et al.Geographical variations in carbon and nitrogen stable isotope ratios in squid[J].Journal of the Marine Biological Association of the United Kingdom,2000,80(4):675-684.
[22] 陈新军,刘必林,王尧耕.世界头足类[M].北京:海洋出版社,2009.CHEN X J,LIU B L,WANG Y G.Cephalopods of the world[M].Beijing:Ocean Press,2009.
[23] DENIRO M J,EPSTEIN S.Influence of diet on the distribution of carbon isotopes in animals[J].Geochimica et Cosmochimica Acta,1978,42(5):495-506.
[24] RUIZ-COOLEY R I,MARKAIDA U,GENDRON D,et al.Stable isotopes in jumbo squid (Dosidicus gigas) beaks to estimate its trophic position:comparison between stomach contents and stable isotopes[J].Journal of the Marine Biological Association of the United Kingdom,2006,86(2):437-445.
[25] NEGRI A,DANERI G A,CEIA F,et al.The cephalopod prey of the Weddell seal,Leptonychotes weddellii,a biological sampler of the Antarctic marine ecosystem[J].Polar Biology,2016,39(3):561-564.
[26] 方舟,金岳,胡飞飞,等.秋季我国近海不同海域短蛸角质颚形态学研究[J].水产学报,2018,42(7):1050-1059.FANG Z,JIN Y,HU F F,et al.Beak morphometrics of short arm octopus (Amphioctopus fangsiao) in different sea areas of China in autumn[J].Journal of Fisheries of China,2018,42(7):1050-1059.
[27] GRAHAM B S,KOCH P L,NEWSOME S D,et al.Using Isoscapes to trace the movements and foraging behavior of top predators in oceanic ecosystems[M]//WEST J B,BOWEN G J,DAWSON T E,et al.Isoscapes:Understanding Movement,Pattern,and Process on Earth through Isotope Mapping.New York:Springer,2010:299-318.
[28] 金岳,陈新军,李云凯,等.基于稳定同位素技术的北太平洋柔鱼角质颚信息[J].生态学杂志,2014,33(8):2101-2107.JIN Y,CHEN X J,LI Y K,et al.Information in beaks of Ommastrephes bartramii in the North Pacific Ocean using stable isotope technology[J].Chinese Journal of Ecology,2014,33(8):2101-2107.
[29] RAU G H,SWEENEY R E,KAPLAN I R.Plankton 13C∶12C ratio changes with latitude:differences between northern and southern oceans[J].Deep Sea Research Part A:Oceanographic Research Papers,1982,29(8):1035-1039.
[30] CHEREL Y,RIDOUX V,SPITZ J,et al.Stable isotopes document the trophic structure of a deep-sea cephalopod assemblage including giant octopod and giant squid[J].Biology Letters,2009,5(3):364-367.
[31] FANG Z,THOMPSON K,JIN Y,et al.Preliminary analysis of beak stable isotopes (δ13C and δ15N) stock variation of neon flying squid,Ommastrephes bartramii,in the North Pacific Ocean[J].Fisheries Research,2016,177:153-163.
[32] LU C C,ICKERINGILL R.Cephalopod beak identification and biomass estimation techniques:tools for dietary studies of southern Australian finfishes[R].Museum Victoria Science Reports.Australia:Fisheries Research and Development Corporation,2002:1-65.