我国异色瓢虫体型大小和色斑类型变异与纬度的关系
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摘要
[目的]体型大小不仅是动物的重要形态特征之一,而且与动物的生活史、生态学和进化密切相关。本研究旨在揭示我国广布的异色瓢虫(Harmonia axyridis)地理种群间形态分化的规律。[方法]从全国纬度在28°(浙江龙泉)至45°(黑龙江哈尔滨)之间的区域采集异色瓢虫样本,测量雌、雄成虫体长及体宽和体积,并观察色斑类型,分析形态特征变异与纬度的关系。[结果]异色瓢虫成虫存在体型性二型现象,雌虫体型显著大于雄虫;雌、雄虫体型大小的差异随纬度升高而减小。雌虫体型不随纬度变化而变化,与贝格曼法则的预测不符;但雄虫体型随纬度上升而增大,符合贝格曼法则。种群中具有淡底色型色斑的异色瓢虫数量占优,具有暗底色型色斑的昆虫较少,暗底色型中黑缘型昆虫极少;具有淡底色型色斑瓢虫占比在高纬度地区略大于低纬度地区。[结论]异色瓢虫体型大小及色斑类型与纬度存在一定相关性。
[Objectives]Animal body size is not only a morphological trait of importance,but also associated with life history,ecology and evolution. This study examined morphological variation among geographical populations of Harmonia axyridis in China.[Methods]Samples of H. axyridis adults were collected at random from the range between 28° and 45° latitudes and measured in their body size( length,width and volume) by sex; their color spot patterns were also categorized. These measures were analyzed in relation to latitude. [Results]Female body was significantly larger than the male,and sexual differences decreased with increasing latitudes. Female body size did not vary with latitude,not in line with the prediction of Bergmann's rule; male body size increased with increasing latitudes,in line with Bergmann's rule. Pale background color spot pattern was overall dominant in populations,followed by the dark background pattern,and then by the black margin pattern. The ladybird beetles with the pale background were more at the high latitude than at the low. [Conclusions]Body size and color spot pattern of H. axyridis vary somewhat with the latitude.
[Objectives]Animal body size is not only a morphological trait of importance,but also associated with life history,ecology and evolution. This study examined morphological variation among geographical populations of Harmonia axyridis in China.[Methods]Samples of H. axyridis adults were collected at random from the range between 28° and 45° latitudes and measured in their body size( length,width and volume) by sex; their color spot patterns were also categorized. These measures were analyzed in relation to latitude. [Results]Female body was significantly larger than the male,and sexual differences decreased with increasing latitudes. Female body size did not vary with latitude,not in line with the prediction of Bergmann's rule; male body size increased with increasing latitudes,in line with Bergmann's rule. Pale background color spot pattern was overall dominant in populations,followed by the dark background pattern,and then by the black margin pattern. The ladybird beetles with the pale background were more at the high latitude than at the low. [Conclusions]Body size and color spot pattern of H. axyridis vary somewhat with the latitude.
引文
[1] Koch R L. The multicolored Asian lady beetle,Harmonia axyridis:a review of its biology,uses in biological control,and non-target impacts[J].Journal of Insect Science,2003,3(1):1-16.
[2] Brown P M,Thomas C E,Lombaert E,et al. The global spread of Harmonia axyridis(Coleoptera:Coccinellidae):distribution,dispersal and routes of invasion[J]. BioControl,2011,56(4):623-641.
[3] Roy H E,Wajnberg E. From biological control to invasion:the ladybird Harmonia axyridis as a model species[J]. BioControl,2008,53(1):1-4.
[4] Roy H E,Brown P M J,Adriaens T,et al. The harlequin ladybird,Harmonia axyridis:global perspectives on invasion history and ecology[J].Biological Invasions,2016,18(4):997-1044.
[5] Vilcinskas A,Stoecker K,Schmidtberg H,et al. Invasive harlequin ladybird carries biological weapons against native competitors[J]. Science,2013,340(6134):862-863.
[6] Lombaert E,Guillemaud T,Lundgren J,et al. Complementarity of statistical treatments to reconstruct worldwide routes of invasion:the case of the Asian ladybird Harmonia axyridis[J]. Molecular Ecology,2014,23(24):5979-5997.
[7] Chown S L,Gaston K J. Body size variation in insects:a macroecological perspective[J]. Biological Reviews,2010,85(1):139-169.
[8] Smith F A,Lyons S K. On being the right size:the impor tance of size in life history,ecology,and evolution[M]//Smith F A,Lyons S K. Animal Body Size:Linking Pattern and Process Across Space,Time,and Taxonomic Group. Chicago:University of Chicago Press,2013:1-12.
[9] Blackburn T M,Gaston K J,Loder N. Geographic gradients in body size:a clarification of Bergmann’s rule[J]. Diversity and Distribution,1999,5(4):165-174.
[10] Blanckenhorn W U,Demont M. Bergmann and converse Bergmann latitudinal clines in arthropods:two ends of a continuum?[J]. Integrative and Comparative Biology,2004,44(6):413-424.
[11] Shelomi M. Where are we now? Bergmann’s rule sensu lato in insects[J]. The American Naturalist,2012,180(4):511-519.
[12] Stillwell R C,Blanckenhorn W U,Teder T,et al. Sex differences in phenotypic plasticity affect variation in sexual size dimorphism in insects:from physiology to evolution[J]. Annual Review of Entomology,2010,55:227-245.
[13] Blanckenhorn W U,Dixon A F,Fairbairn D J,et al. Proximate causes of Rensch’s rule:does sexual size dimorphism in arthropods result from sex differences in development time?[J]. The American Naturalist,2007,169(2):245-257.
[14] Gaston K J,Chown S L. Macroecological patterns in insect body size[M]//Smith F A,Lyons S K. Animal Body Size:Linking Pattern and Process Across Space,Time,and Taxonomic Group. Chicago:University of Chicago Press,2013:13-61.
[15] Majerus M E. A Natural History of Ladybird Beetles[M]. Cambridge:Cambridge University Press,2016.
[16] Kawakami Y,Yamazaki K,Ohashi K. Geographical variations of elytral color polymorphism in Cheilomenes sexmaculata(Fabricius)(Coleoptera:Coccinellidae)[J]. Entomological Science,2013,16(2):235-242.
[17]郭长飞,许炜明,何瞻,等.异色瓢虫色斑多样性调查及各地十九斑变型COI基因系统进化分析[J].植物保护学报,2016,43(1):155-161.Guo C F,Xu W M,He Z,et al. Color variations of Harmonia axyridis and phylogenetic analysis of 19-spot H. axyridis based on their COI genes[J].Journal of Plant Protection,2016,43(1):155-161(in Chinese with English abstract).
[18]宫庆涛,耿军,张坤鹏,等.泰安市异色瓢虫色斑型和性比变化[J].环境昆虫学报,2016,38(2):313-323.Gong Q T,Geng J,Zhang K P,et al. Phenotypes and sex ratio changes of Harmonia axyridis(Pallas)in Tai’an[J]. Journal of Environmental Entomology,2016,38(2):313-323(in Chinese with English abstract).
[19] Li J,Ke X,Wang Z,et al. Investigation on color pattern diversity of overwintering Harmonia axyridis in southern suburb of Jiamusi[J]. Plant Diseases and Pests,2011,2(4):40-42.
[20] Osawa N,Nishida T. Seasonal variation in elytral colour polymorphism in Harmonia axyridis(the ladybird beetle):the role of non-random mating[J].Heredity,1992,69(4):297-307.
[21] Wang S,Michaud J P,Tan X L,et al. Melanism in a Chinese population of Harmonia axyridis(Coleoptera:Coccinellidae):a criterion for male investment with pleiotropic effects on behavior and fertility[J]. Journal of Insect Behavior,2013,26(5):679-689.
[22] Ueno H,Sato Y,Tsuchida K. Colour-associated mating success in a polymorphic ladybird beetle,Harmonia axyridis[J]. Functional Ecology,1998,12(5):757-761.
[23] Atkinson W D. A comparison of the reproductive strategies of domestic species of Drosophila[J]. Journal of Animal Ecology,1979,48(1):53-64.
[24] Lovich J E,Gibbons J W. A review of techniques for quantifying sexual size dimorphism[J]. Growth Development and Aging,1992,56:269-281.
[25] R Core Team. R:A Language and Environment for Statistical Computing[EB/OL]. R Foundation for Statistical Computing,Vienna,Austria.[2016-10-31]. https://www.r-project.org/.
[26] Fairbairn D J,Blanckenhorn W U,Székely T. Sex,Size and Gender Roles:Evolutionary Studies of Sexual Size Dimorphism[M]. London:Oxford University Press,2007.
[27] García-Navas V,Noguerales V,Cordero P J,et al. Ecological drivers of body size evolution and sexual size dimorphism in short-horned grasshoppers(Orthoptera:Acrididae)[J]. Journal of Evolutionary Biology,2017,30(8):1592-1608.
[28] Cirino L A,Miller C W. Seasonal effects on the population,morphology and reproductive behavior of Narnia femorata(Hemiptera:Coreidae)[J].Insects,2017,8(1):13.
[29] Stillwell R C,Fox C W. Environmental effects on sexual size dimorphism of a seed-feeding beetle[J]. Oecologia,2007,153(2):273-280.
[30] Gianoli E,Suarez L H,Gonzales W L,et al. Host-associated variation in sexual size dimorphism and fitness effects of adult feeding in a bruchid beetle[J]. Entomologia Experimentalis et Applicata,2007,122(3):233-237.
[31] Stillwell R C,Morse G E,Fox C W. Geographic variation in body size and sexual size dimorphism of a seed-feeding beetle[J]. The American Naturalist,2007,170(3):358-369.
[32]毕孟杰,沈梦伟,周可新,等.中国瓢虫体型的地理分异及其与环境因子的关系[J].生物多样性,2015,23(6):775-783.Bi M J,Shen M W,Zhou K X,et al. Geographical variance of ladybird morphology and environmental correlates in China[J]. Biodiversity Science,2015,23(6):775-783(in Chinese with English abstract).
[33] Berger D,Walters R,Gotthard K. What limits insect fecundity? Body size-and temperature-dependent egg maturation and oviposition in a butterfly[J]. Functional Ecology,2008,22(3):523-529.
[34] Honěk A. Intraspecific variation in body size and fecundity in insects:a general relationship[J]. Oikos,1993,66(3):483-492.
[35] Py2ek P,Richardson D M. Invasive species,environmental change and management,and health[J]. Annual Review of Environment and Resources,2010,35:25-55.
[36]赵天璇,袁明龙.我国异色瓢虫的生物生态学特性[J].草业科学,2017,34(3):614-629.Zhao T X,Yuan M L. Biological and ecological characteristics of Harmonia axyridis in China[J]. Pratacultural Science,2017,34(3):614-629(in Chinese with English abstract).
[37]杜文梅,陈丽玲,阮长春,等.长春市异色瓢虫色斑类型调查与研究[J].中国农学通报,2010,26(4):258-266.Du W M,Chen L L,Ruan C C,et al. Investigations of the color variations of Harmonia axyridis(Pallas)in Changchun[J]. Chinese Agricultural Science Bulletin,2010,26(4):258-266(in Chinese with English abstract).
[38]徐青叶,林青青,金上,等.临安大明山异色瓢虫及龟纹瓢虫鞘翅色斑多样性分析[J].杭州师范大学学报(自然科学版),2014,13(2):159-163.Xu Q Y,Lin Q Q,Jin S,et al. On the elytra colored speck diversity of Harmonia axyridis(Pallas)and Propylea japonica(Thunberg)in Daming mountain[J]. Journal of Hangzhou Normal University(Natural Science Edition),2014,13(2):159-163(in Chinese with English abstract).
[39] Wang S,Michaud J P,Zhang R,et al. Seasonal cycles of assortative mating and reproductive behaviour in polymorphic populations of Harmonia axyridis in China[J]. Ecological Entomology,2009,34(4):483-494.
[40] Dobzhansky T. Geographical variation in lady-beetles[J]. The American Naturalist,1933,67(709):97-126.
[2] Brown P M,Thomas C E,Lombaert E,et al. The global spread of Harmonia axyridis(Coleoptera:Coccinellidae):distribution,dispersal and routes of invasion[J]. BioControl,2011,56(4):623-641.
[3] Roy H E,Wajnberg E. From biological control to invasion:the ladybird Harmonia axyridis as a model species[J]. BioControl,2008,53(1):1-4.
[4] Roy H E,Brown P M J,Adriaens T,et al. The harlequin ladybird,Harmonia axyridis:global perspectives on invasion history and ecology[J].Biological Invasions,2016,18(4):997-1044.
[5] Vilcinskas A,Stoecker K,Schmidtberg H,et al. Invasive harlequin ladybird carries biological weapons against native competitors[J]. Science,2013,340(6134):862-863.
[6] Lombaert E,Guillemaud T,Lundgren J,et al. Complementarity of statistical treatments to reconstruct worldwide routes of invasion:the case of the Asian ladybird Harmonia axyridis[J]. Molecular Ecology,2014,23(24):5979-5997.
[7] Chown S L,Gaston K J. Body size variation in insects:a macroecological perspective[J]. Biological Reviews,2010,85(1):139-169.
[8] Smith F A,Lyons S K. On being the right size:the impor tance of size in life history,ecology,and evolution[M]//Smith F A,Lyons S K. Animal Body Size:Linking Pattern and Process Across Space,Time,and Taxonomic Group. Chicago:University of Chicago Press,2013:1-12.
[9] Blackburn T M,Gaston K J,Loder N. Geographic gradients in body size:a clarification of Bergmann’s rule[J]. Diversity and Distribution,1999,5(4):165-174.
[10] Blanckenhorn W U,Demont M. Bergmann and converse Bergmann latitudinal clines in arthropods:two ends of a continuum?[J]. Integrative and Comparative Biology,2004,44(6):413-424.
[11] Shelomi M. Where are we now? Bergmann’s rule sensu lato in insects[J]. The American Naturalist,2012,180(4):511-519.
[12] Stillwell R C,Blanckenhorn W U,Teder T,et al. Sex differences in phenotypic plasticity affect variation in sexual size dimorphism in insects:from physiology to evolution[J]. Annual Review of Entomology,2010,55:227-245.
[13] Blanckenhorn W U,Dixon A F,Fairbairn D J,et al. Proximate causes of Rensch’s rule:does sexual size dimorphism in arthropods result from sex differences in development time?[J]. The American Naturalist,2007,169(2):245-257.
[14] Gaston K J,Chown S L. Macroecological patterns in insect body size[M]//Smith F A,Lyons S K. Animal Body Size:Linking Pattern and Process Across Space,Time,and Taxonomic Group. Chicago:University of Chicago Press,2013:13-61.
[15] Majerus M E. A Natural History of Ladybird Beetles[M]. Cambridge:Cambridge University Press,2016.
[16] Kawakami Y,Yamazaki K,Ohashi K. Geographical variations of elytral color polymorphism in Cheilomenes sexmaculata(Fabricius)(Coleoptera:Coccinellidae)[J]. Entomological Science,2013,16(2):235-242.
[17]郭长飞,许炜明,何瞻,等.异色瓢虫色斑多样性调查及各地十九斑变型COI基因系统进化分析[J].植物保护学报,2016,43(1):155-161.Guo C F,Xu W M,He Z,et al. Color variations of Harmonia axyridis and phylogenetic analysis of 19-spot H. axyridis based on their COI genes[J].Journal of Plant Protection,2016,43(1):155-161(in Chinese with English abstract).
[18]宫庆涛,耿军,张坤鹏,等.泰安市异色瓢虫色斑型和性比变化[J].环境昆虫学报,2016,38(2):313-323.Gong Q T,Geng J,Zhang K P,et al. Phenotypes and sex ratio changes of Harmonia axyridis(Pallas)in Tai’an[J]. Journal of Environmental Entomology,2016,38(2):313-323(in Chinese with English abstract).
[19] Li J,Ke X,Wang Z,et al. Investigation on color pattern diversity of overwintering Harmonia axyridis in southern suburb of Jiamusi[J]. Plant Diseases and Pests,2011,2(4):40-42.
[20] Osawa N,Nishida T. Seasonal variation in elytral colour polymorphism in Harmonia axyridis(the ladybird beetle):the role of non-random mating[J].Heredity,1992,69(4):297-307.
[21] Wang S,Michaud J P,Tan X L,et al. Melanism in a Chinese population of Harmonia axyridis(Coleoptera:Coccinellidae):a criterion for male investment with pleiotropic effects on behavior and fertility[J]. Journal of Insect Behavior,2013,26(5):679-689.
[22] Ueno H,Sato Y,Tsuchida K. Colour-associated mating success in a polymorphic ladybird beetle,Harmonia axyridis[J]. Functional Ecology,1998,12(5):757-761.
[23] Atkinson W D. A comparison of the reproductive strategies of domestic species of Drosophila[J]. Journal of Animal Ecology,1979,48(1):53-64.
[24] Lovich J E,Gibbons J W. A review of techniques for quantifying sexual size dimorphism[J]. Growth Development and Aging,1992,56:269-281.
[25] R Core Team. R:A Language and Environment for Statistical Computing[EB/OL]. R Foundation for Statistical Computing,Vienna,Austria.[2016-10-31]. https://www.r-project.org/.
[26] Fairbairn D J,Blanckenhorn W U,Székely T. Sex,Size and Gender Roles:Evolutionary Studies of Sexual Size Dimorphism[M]. London:Oxford University Press,2007.
[27] García-Navas V,Noguerales V,Cordero P J,et al. Ecological drivers of body size evolution and sexual size dimorphism in short-horned grasshoppers(Orthoptera:Acrididae)[J]. Journal of Evolutionary Biology,2017,30(8):1592-1608.
[28] Cirino L A,Miller C W. Seasonal effects on the population,morphology and reproductive behavior of Narnia femorata(Hemiptera:Coreidae)[J].Insects,2017,8(1):13.
[29] Stillwell R C,Fox C W. Environmental effects on sexual size dimorphism of a seed-feeding beetle[J]. Oecologia,2007,153(2):273-280.
[30] Gianoli E,Suarez L H,Gonzales W L,et al. Host-associated variation in sexual size dimorphism and fitness effects of adult feeding in a bruchid beetle[J]. Entomologia Experimentalis et Applicata,2007,122(3):233-237.
[31] Stillwell R C,Morse G E,Fox C W. Geographic variation in body size and sexual size dimorphism of a seed-feeding beetle[J]. The American Naturalist,2007,170(3):358-369.
[32]毕孟杰,沈梦伟,周可新,等.中国瓢虫体型的地理分异及其与环境因子的关系[J].生物多样性,2015,23(6):775-783.Bi M J,Shen M W,Zhou K X,et al. Geographical variance of ladybird morphology and environmental correlates in China[J]. Biodiversity Science,2015,23(6):775-783(in Chinese with English abstract).
[33] Berger D,Walters R,Gotthard K. What limits insect fecundity? Body size-and temperature-dependent egg maturation and oviposition in a butterfly[J]. Functional Ecology,2008,22(3):523-529.
[34] Honěk A. Intraspecific variation in body size and fecundity in insects:a general relationship[J]. Oikos,1993,66(3):483-492.
[35] Py2ek P,Richardson D M. Invasive species,environmental change and management,and health[J]. Annual Review of Environment and Resources,2010,35:25-55.
[36]赵天璇,袁明龙.我国异色瓢虫的生物生态学特性[J].草业科学,2017,34(3):614-629.Zhao T X,Yuan M L. Biological and ecological characteristics of Harmonia axyridis in China[J]. Pratacultural Science,2017,34(3):614-629(in Chinese with English abstract).
[37]杜文梅,陈丽玲,阮长春,等.长春市异色瓢虫色斑类型调查与研究[J].中国农学通报,2010,26(4):258-266.Du W M,Chen L L,Ruan C C,et al. Investigations of the color variations of Harmonia axyridis(Pallas)in Changchun[J]. Chinese Agricultural Science Bulletin,2010,26(4):258-266(in Chinese with English abstract).
[38]徐青叶,林青青,金上,等.临安大明山异色瓢虫及龟纹瓢虫鞘翅色斑多样性分析[J].杭州师范大学学报(自然科学版),2014,13(2):159-163.Xu Q Y,Lin Q Q,Jin S,et al. On the elytra colored speck diversity of Harmonia axyridis(Pallas)and Propylea japonica(Thunberg)in Daming mountain[J]. Journal of Hangzhou Normal University(Natural Science Edition),2014,13(2):159-163(in Chinese with English abstract).
[39] Wang S,Michaud J P,Zhang R,et al. Seasonal cycles of assortative mating and reproductive behaviour in polymorphic populations of Harmonia axyridis in China[J]. Ecological Entomology,2009,34(4):483-494.
[40] Dobzhansky T. Geographical variation in lady-beetles[J]. The American Naturalist,1933,67(709):97-126.