南海鸢乌贼的遗传差异:种群分化还是种间分化
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摘要
利用线粒体ND2、COI和16SrDNA序列,对2012年9月至2015年9月采自南海的鸢乌贼(Sthenoteuthis oualaniensis)样本进行了种群遗传结构的分析。系统发育关系显示鸢乌贼中型群和微型群各自形成了单系群分支,而分支内没有形成与地理群体或季节群体对应的支系。分子方差分析同样表明,地理群体或季节群体之间不存在明显的遗传分化,而中型群和微型群之间具有极显著的遗传分化(F_(ST)=0.94092～0.9899, P<0.0001)。遗传多样性分析显示,鸢乌贼中型群和微型群均呈现出较高的单倍型多样性(0.9465和0.8545)和较低的核苷酸多样性(0.0051和0.0021),中型群的遗传多样性指数要高于微型群。基于ND2、COI和16SrDNA序列计算的中型群和微型群之间的遗传距离分别达到了14.0%、9.6%和8.8%,远大于类群内的遗传分化(平均<1%)。研究结果表明,南海鸢乌贼存在分布高度重合且有显著遗传分化的两个种群,即中型群和微型群。中型群和微型群之间的遗传差异达到了种间分化的水平,提示体型较大、具有背部发光器的中型群鸢乌贼(典型鸢乌贼)与体型较小、背部发光器缺失的微型群鸢乌贼可能是两个相互独立的种类。
In the present study, the population differentiation of the purpleback flying squid(Sthenoteuthis oualaniensis) in the South China Sea was examined based on mitochondrial ND2, COI, and 16 S rDNA sequences, using samples collected between September 2012 and September 2015. Phylogenetic analysis revealed two significantly divergent monophyletic groups corresponding to the "medium form" and the "dwarf form, " but no geographical lineages corresponding to the sampling localities or seasonal groups were found. The Analysis of Molecular Variance also revealed significant genetic differentiation between the "medium form" and the "dwarf form"(F_(ST)=0.94092-0.9899, P<0.0001), while no genetic variance between either geographic populations or seasonal groups was found. The genetic diversity analysis showed that both the "medium form" and the "dwarf form" of S. oualaniensis in the South China Sea were characterized by high haplotype diversity(0.9465 and 0.8545) and low nucleotide diversity(0.0051 and 0.0021). The genetic distance between the ‘medium form' and the "dwarf form" based on ND2, COI, and 16 S sequences were 14.0%, 9.6%, and 8.8%, respectively. The genetic distances between the two forms were considerably larger than the genetic distances within each of the two forms(<1%). It was demonstrated that two divergent populations of S. oualaniensis were sympatrically distributed in the South China Sea, i.e. the "medium form" and the "dwarf form." The genetic differentiation between these two populations was similar to that among separate species within Ommastrephidae, which indicated that the two morphologically distinct forms of the purpleback flying squid, namely the "medium form"(middle sized, with a photophore) and the "dwarf form"(small sized, without a photophore), are two distinct species.
In the present study, the population differentiation of the purpleback flying squid(Sthenoteuthis oualaniensis) in the South China Sea was examined based on mitochondrial ND2, COI, and 16 S rDNA sequences, using samples collected between September 2012 and September 2015. Phylogenetic analysis revealed two significantly divergent monophyletic groups corresponding to the "medium form" and the "dwarf form, " but no geographical lineages corresponding to the sampling localities or seasonal groups were found. The Analysis of Molecular Variance also revealed significant genetic differentiation between the "medium form" and the "dwarf form"(F_(ST)=0.94092-0.9899, P<0.0001), while no genetic variance between either geographic populations or seasonal groups was found. The genetic diversity analysis showed that both the "medium form" and the "dwarf form" of S. oualaniensis in the South China Sea were characterized by high haplotype diversity(0.9465 and 0.8545) and low nucleotide diversity(0.0051 and 0.0021). The genetic distance between the ‘medium form' and the "dwarf form" based on ND2, COI, and 16 S sequences were 14.0%, 9.6%, and 8.8%, respectively. The genetic distances between the two forms were considerably larger than the genetic distances within each of the two forms(<1%). It was demonstrated that two divergent populations of S. oualaniensis were sympatrically distributed in the South China Sea, i.e. the "medium form" and the "dwarf form." The genetic differentiation between these two populations was similar to that among separate species within Ommastrephidae, which indicated that the two morphologically distinct forms of the purpleback flying squid, namely the "medium form"(middle sized, with a photophore) and the "dwarf form"(small sized, without a photophore), are two distinct species.
引文
[1]Dunning M C.A review of the systematics,distribution,and biology of the arrow squid genera Sthenoteuthis Verrill,1880,and Ornithoteuthis Okada,1927(Cephalopoda:Ommastrephidae)[J].Smithson Contribution Zoology,1998,586(2):425-433.
[2]Chen X J,Liu B L,Wang Y G.Cephalopods of the World[M].Beijing:China Ocean Press,2009:312-313.[陈新军,刘必林,王尧耕.世界头足类[M].北京:海洋出版社,2009:312-313.]
[3]Wang Y G,Chen X J.The World Ocean Ecomomic Squid Resources and Fisheries[M].Beijing:China Ocean Press,2005:284-295.[王尧耕,陈新军.世界大洋性经济柔鱼类资源及其渔业[M].北京:海洋出版社,2005:284-295.]
[4]Zhang P,Yang L,Zhang X F,et al.The present status and prospect on exploitation of tuna and squid fishery resources in South China Sea[J].South China Fisheries Science,2010,6(1):68-74.[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.]
[5]Chen G B,Zhang J,Yu J,et al.Hydroacoustic scattering characteristics and biomass assessment of the purpleback flying squid[Sthenoteuthis oualaniensis,(Lesson,1830)]from the deep water area of the South China Sea[J].Journal of Applied Ichthyology,2013,29(6):1447-1452.
[6]Zhang J,Chen G B,Zhang P,et al.Estimation of purpleback flying quid(Sthenoteuthis oualaniensis)resource in the central and southern South China Sea based on fisheries acoustics and light-falling net[J].Journal of Fishery Sciences of China,2014,21(4):822-831.[张俊,陈国宝,张鹏,等.基于渔业声学和灯光罩网的南海中南部鸢乌贼资源评估[J].中国水产科学,2014,21(4):822-831.]
[7]Nesis K N.Population structure of oceanic ommastrephids,with particular reference to Sthenoteuthis oualaniensis:A review[M]//Recent Advances in Fisheries Biology.Tokyo:Tokai University Press,1993:375-383.
[8]Staaf D J,Ruiz-Cooley R I,Elliger C,et al.Ommastrephid squids Sthenoteuthis oualaniensis and Dosidicus gigas in the eastern Pacific show convergent biogeographic breaks but contrasting population structures[J].Marine Ecology Progress Series,2010,418:165-178.
[9]Yan Y R,Feng B,Lu H S,et al.Fishery biology of purpleback flying squid Sthenoteuthis oualaniensis in northern sea areas around Nansha islands in summer[J].Oceanologia et Limnologia Sinica,2012,43(6):1177-1186.[颜云榕,冯波,卢伙胜,等.南沙群岛北部海域鸢乌贼(Sthenoteuthis oualaniensis)夏季渔业生物学研究[J].海洋与湖沼,2012.43(6):1177-1186.]
[10]Zhang P,Yan L,Yang B Z,et al.Population structure of purpleback flying squid(Sthenoteuthis oualaniensis)in Nansha area in spring[J].South China Fisheries Science,2015,11(5):11-19.[张鹏,晏磊,杨炳忠,等.春季南沙海域鸢乌贼种群结构特征的研究[J].南方水产科学,2015,11(5):11-19.]
[11]Fan J T,Qiu Y S,Chen Z Z,et al.Morphological difference of the beak between two stocks of Sthenoteuthis oualaniensis inhabiting South China Sea[J].Periodical of Ocean University of China,2015,45(10):42-49.[范江涛,邱永松,陈作志,等.南海鸢乌贼两个群体间角质颚形态差异分析[J].中国海洋大学学报,2015,45(10):42-49.]
[12]Zhu K,Wang X H,Zhang P,et al.A study on morphological variations and discrimination of medium and dwarf forms of purple flying squid Sthenoteuthis oualaniensis in the southern South China Sea[J].Journal of Tropical Oceanography,2016,35(6):82-88.[朱凯,王雪辉,张鹏,等,南海南部鸢乌贼中型群与微型群形态学差异及其判别分析[J].热带海洋学报,2016,35(6):82-88.]
[13]Yan Y R,Yi M R,Feng B,et al.Morphological variations and discriminant analysis of three different geographic cohorts of Sthenoteuthis oualaniensis in the South China Sea[J].Journal of Guangdong Ocean University,2015,35(3):43-50.[颜云榕,易木荣,冯波,等.南海鸢乌贼3个地理群体形态差异与判别分析[J].广东海洋大学学报,2015,35(3):43-50.]
[14]Yang D K.The resources and its exploitation and utilization of two species of squid[J].Journal of Shanghai Fisheries University,2002,11(2):176-179.[杨德康.两种鱿鱼资源和其开发利用[J].上海水产大学学报,2002,11(2):176-179.]
[15]Okutani T,Tung I.Reviews of biology of commercially important squids in Japanese and adjacent waters.I.Symplectoteuthis oualaniensis(Lesson)[J].Veliger,1978,21(1):87-94.
[16]Hall T A.BioEdit:a user-friendly biological sequence alignment editor and analysis program for Windows95/98/NT[J].Nucleic Acids Symposium Series,1999,41:95-98.
[17]Kumar S,Stecher G,Tamura K.MEGA7:Molecular evolutionary genetics analysis version 7.0 for bigger datasets[J].Molecular Biology Evolution,2016,33(7):1870-1874.
[18]Edgar R C.MUSCLE:multiple sequence alignment with high accuracy and high throughput[J].Nucleic Acids Research,2004,32(5):1792-1797.
[19]Tamura K,Nei M.Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees[J].Molecular Biology Evolution,1993,10(3):512-526.
[20]Tamura K.Estimation of the number of nucleotide substitutions when there are strong transition-transversion and G+C-content biases[J].Molecular Biology Evolution,1992,9(4):678-687.
[21]Excoffier L,Lischer H L.Arlequin suite ver 3.5:a new series of programs to perform population genetics analyses under Linux and Windows[J].Molecular Ecology Resource,2010,10(3):564-567.
[22]Excoffier L,Smouse P E,Quattro J M.Analysis of molecular variance inferred from metric distances among DNAhaplotypes:application to human mitochondrial DNA restriction data[J].Genetics,1992,131(2):479-491.
[23]Librado P,Rozas J.DnaSP v5:a software for comprehensive analysis of DNA polymorphism data[J].Bioinformatics,2009,25(11):1451-1452.
[24]Roldán M I,Planella L,Heras S,et al.Genetic analyses of two spawning stocks of the short-finned squid(Illex argentinus)using nuclear and mitochondrial data[J].Comptes Rendus Biologies,2014,337(9):503-512.
[25]Freeland J R.Molecular Ecology[M].Chichester:John Wiley&Sons Ltd.,2005:112-116.
[26]Wright S.The genetical structure of populations[J].Annals of Human Genetics,1949,15(1):323-354.
[27]Wright S.Evolution and the Genetics of Population:Variability within and among Natural Population[M].Chicago:University of Chicago Press,1978:79-103.
[28]Wormuth J H.The Biogeography and Numerical Taxonomy of the Oegopsid Squid Family Ommastrephidae in the Pacific Ocean[M].California:University of California Press,1976.
[29]Young R E.A brief review of the biology of the oceanic squid,Symplectoteuthis oualaniensis(Lesson)[J].Comparative Biochemistry and Physiology Part B:Comparative Biochemistry,1975,52(1):141-143.
[30]Young R E,Michael V.Sthenoteuthis oualaniensis(Lesson,1830).Version 10 November 2012(under construction)[Z].http://tolweb.org/Sthenoteuthis_oualaniensis/77447.
[31]Snyder R.Aspects of the biology of the giant form of Sthenoteuthis oualaniensis(Cephalopoda:Ommastrephidae)from the Arabian Sea[J].Journal of Molluscan Studies,1998,64(1):21-34.
[32]Clarke M R.Large light organs on the dorsal surface of the squids Ommastrephes pteropus,‘Symplectoteuthis oualaniensis’and‘Dosidicus gigas’[J].Journal of Molluscan Studies,1965,36(5):319-321.
[33]Hebert P D N,Cywinska A,Ball S L.Biological identifications through DNA barcodes[J].Proceedings of the Royal Society of London B:Biological Sciences,2003,270(1512):313-321.
[34]Yu H,Kong L,Li Q.Evaluation of the efficacy of twelve mitochondrial protein-coding genes as barcodes for mollusk DNA barcoding[J].Mitochondrial DNA Part A,2016,27(2):1336-1339.
[35]Carvalho G R,Thompson A,Stoner A L.Genetic diversity and population differentiation of the shortfin squid Illex argentinus in the south-west Atlantic[J].Journal of Experimental Marine Biology and Ecology,1992,158(1):105-121.
[36]Nesis K.Population structure of the squid Sthenoteuthis oualaniensis(Lesson,1830)in the tropical West Pacific[J].Trudy Instituta Okeanologii Akademiia Nauk SSSR,1977,107:15-29.
[37]Roeleveld M A C,Lipinski M.The giant squid Architeuthis in southern African waters[J].Journal of Zoology,2009,224(3):431-477.
[2]Chen X J,Liu B L,Wang Y G.Cephalopods of the World[M].Beijing:China Ocean Press,2009:312-313.[陈新军,刘必林,王尧耕.世界头足类[M].北京:海洋出版社,2009:312-313.]
[3]Wang Y G,Chen X J.The World Ocean Ecomomic Squid Resources and Fisheries[M].Beijing:China Ocean Press,2005:284-295.[王尧耕,陈新军.世界大洋性经济柔鱼类资源及其渔业[M].北京:海洋出版社,2005:284-295.]
[4]Zhang P,Yang L,Zhang X F,et al.The present status and prospect on exploitation of tuna and squid fishery resources in South China Sea[J].South China Fisheries Science,2010,6(1):68-74.[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.]
[5]Chen G B,Zhang J,Yu J,et al.Hydroacoustic scattering characteristics and biomass assessment of the purpleback flying squid[Sthenoteuthis oualaniensis,(Lesson,1830)]from the deep water area of the South China Sea[J].Journal of Applied Ichthyology,2013,29(6):1447-1452.
[6]Zhang J,Chen G B,Zhang P,et al.Estimation of purpleback flying quid(Sthenoteuthis oualaniensis)resource in the central and southern South China Sea based on fisheries acoustics and light-falling net[J].Journal of Fishery Sciences of China,2014,21(4):822-831.[张俊,陈国宝,张鹏,等.基于渔业声学和灯光罩网的南海中南部鸢乌贼资源评估[J].中国水产科学,2014,21(4):822-831.]
[7]Nesis K N.Population structure of oceanic ommastrephids,with particular reference to Sthenoteuthis oualaniensis:A review[M]//Recent Advances in Fisheries Biology.Tokyo:Tokai University Press,1993:375-383.
[8]Staaf D J,Ruiz-Cooley R I,Elliger C,et al.Ommastrephid squids Sthenoteuthis oualaniensis and Dosidicus gigas in the eastern Pacific show convergent biogeographic breaks but contrasting population structures[J].Marine Ecology Progress Series,2010,418:165-178.
[9]Yan Y R,Feng B,Lu H S,et al.Fishery biology of purpleback flying squid Sthenoteuthis oualaniensis in northern sea areas around Nansha islands in summer[J].Oceanologia et Limnologia Sinica,2012,43(6):1177-1186.[颜云榕,冯波,卢伙胜,等.南沙群岛北部海域鸢乌贼(Sthenoteuthis oualaniensis)夏季渔业生物学研究[J].海洋与湖沼,2012.43(6):1177-1186.]
[10]Zhang P,Yan L,Yang B Z,et al.Population structure of purpleback flying squid(Sthenoteuthis oualaniensis)in Nansha area in spring[J].South China Fisheries Science,2015,11(5):11-19.[张鹏,晏磊,杨炳忠,等.春季南沙海域鸢乌贼种群结构特征的研究[J].南方水产科学,2015,11(5):11-19.]
[11]Fan J T,Qiu Y S,Chen Z Z,et al.Morphological difference of the beak between two stocks of Sthenoteuthis oualaniensis inhabiting South China Sea[J].Periodical of Ocean University of China,2015,45(10):42-49.[范江涛,邱永松,陈作志,等.南海鸢乌贼两个群体间角质颚形态差异分析[J].中国海洋大学学报,2015,45(10):42-49.]
[12]Zhu K,Wang X H,Zhang P,et al.A study on morphological variations and discrimination of medium and dwarf forms of purple flying squid Sthenoteuthis oualaniensis in the southern South China Sea[J].Journal of Tropical Oceanography,2016,35(6):82-88.[朱凯,王雪辉,张鹏,等,南海南部鸢乌贼中型群与微型群形态学差异及其判别分析[J].热带海洋学报,2016,35(6):82-88.]
[13]Yan Y R,Yi M R,Feng B,et al.Morphological variations and discriminant analysis of three different geographic cohorts of Sthenoteuthis oualaniensis in the South China Sea[J].Journal of Guangdong Ocean University,2015,35(3):43-50.[颜云榕,易木荣,冯波,等.南海鸢乌贼3个地理群体形态差异与判别分析[J].广东海洋大学学报,2015,35(3):43-50.]
[14]Yang D K.The resources and its exploitation and utilization of two species of squid[J].Journal of Shanghai Fisheries University,2002,11(2):176-179.[杨德康.两种鱿鱼资源和其开发利用[J].上海水产大学学报,2002,11(2):176-179.]
[15]Okutani T,Tung I.Reviews of biology of commercially important squids in Japanese and adjacent waters.I.Symplectoteuthis oualaniensis(Lesson)[J].Veliger,1978,21(1):87-94.
[16]Hall T A.BioEdit:a user-friendly biological sequence alignment editor and analysis program for Windows95/98/NT[J].Nucleic Acids Symposium Series,1999,41:95-98.
[17]Kumar S,Stecher G,Tamura K.MEGA7:Molecular evolutionary genetics analysis version 7.0 for bigger datasets[J].Molecular Biology Evolution,2016,33(7):1870-1874.
[18]Edgar R C.MUSCLE:multiple sequence alignment with high accuracy and high throughput[J].Nucleic Acids Research,2004,32(5):1792-1797.
[19]Tamura K,Nei M.Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees[J].Molecular Biology Evolution,1993,10(3):512-526.
[20]Tamura K.Estimation of the number of nucleotide substitutions when there are strong transition-transversion and G+C-content biases[J].Molecular Biology Evolution,1992,9(4):678-687.
[21]Excoffier L,Lischer H L.Arlequin suite ver 3.5:a new series of programs to perform population genetics analyses under Linux and Windows[J].Molecular Ecology Resource,2010,10(3):564-567.
[22]Excoffier L,Smouse P E,Quattro J M.Analysis of molecular variance inferred from metric distances among DNAhaplotypes:application to human mitochondrial DNA restriction data[J].Genetics,1992,131(2):479-491.
[23]Librado P,Rozas J.DnaSP v5:a software for comprehensive analysis of DNA polymorphism data[J].Bioinformatics,2009,25(11):1451-1452.
[24]Roldán M I,Planella L,Heras S,et al.Genetic analyses of two spawning stocks of the short-finned squid(Illex argentinus)using nuclear and mitochondrial data[J].Comptes Rendus Biologies,2014,337(9):503-512.
[25]Freeland J R.Molecular Ecology[M].Chichester:John Wiley&Sons Ltd.,2005:112-116.
[26]Wright S.The genetical structure of populations[J].Annals of Human Genetics,1949,15(1):323-354.
[27]Wright S.Evolution and the Genetics of Population:Variability within and among Natural Population[M].Chicago:University of Chicago Press,1978:79-103.
[28]Wormuth J H.The Biogeography and Numerical Taxonomy of the Oegopsid Squid Family Ommastrephidae in the Pacific Ocean[M].California:University of California Press,1976.
[29]Young R E.A brief review of the biology of the oceanic squid,Symplectoteuthis oualaniensis(Lesson)[J].Comparative Biochemistry and Physiology Part B:Comparative Biochemistry,1975,52(1):141-143.
[30]Young R E,Michael V.Sthenoteuthis oualaniensis(Lesson,1830).Version 10 November 2012(under construction)[Z].http://tolweb.org/Sthenoteuthis_oualaniensis/77447.
[31]Snyder R.Aspects of the biology of the giant form of Sthenoteuthis oualaniensis(Cephalopoda:Ommastrephidae)from the Arabian Sea[J].Journal of Molluscan Studies,1998,64(1):21-34.
[32]Clarke M R.Large light organs on the dorsal surface of the squids Ommastrephes pteropus,‘Symplectoteuthis oualaniensis’and‘Dosidicus gigas’[J].Journal of Molluscan Studies,1965,36(5):319-321.
[33]Hebert P D N,Cywinska A,Ball S L.Biological identifications through DNA barcodes[J].Proceedings of the Royal Society of London B:Biological Sciences,2003,270(1512):313-321.
[34]Yu H,Kong L,Li Q.Evaluation of the efficacy of twelve mitochondrial protein-coding genes as barcodes for mollusk DNA barcoding[J].Mitochondrial DNA Part A,2016,27(2):1336-1339.
[35]Carvalho G R,Thompson A,Stoner A L.Genetic diversity and population differentiation of the shortfin squid Illex argentinus in the south-west Atlantic[J].Journal of Experimental Marine Biology and Ecology,1992,158(1):105-121.
[36]Nesis K.Population structure of the squid Sthenoteuthis oualaniensis(Lesson,1830)in the tropical West Pacific[J].Trudy Instituta Okeanologii Akademiia Nauk SSSR,1977,107:15-29.
[37]Roeleveld M A C,Lipinski M.The giant squid Architeuthis in southern African waters[J].Journal of Zoology,2009,224(3):431-477.