珠江水系鱼类群落多样性空间分布格局
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摘要
珠江是我国南方第一大河,是我国重要淡水渔业生产基地和水生生物资源基因库。珠江鱼类在维持生物多样性、提供鱼类种质资源方面举足轻重。但是到目前为止,关于其鱼类空间分布格局的研究甚少。特别是近几十年来各种水工建设和过度捕捞使得渔业资源急剧衰退,鱼类空间分布的研究显得尤为重要。2015年对珠江全流域13个站位进行了全面调查,共采集渔获物10119尾,隶属于94种72属17科。鲤科鱼类占显著优势,其次种类较多的依次为鲿科、鳅科。采用非度量多维标度排序(NMDS)方法对鱼类群落空间分布特征进行了分析,结果表明珠江鱼类被划分为3个类群,即以餐、南方拟餐、黄颡鱼等小型鱼类为主的中上游类群、以赤眼鳟、鲮鱼、广东鲂等中型鱼类为主的中下游类群和以罗非鱼为主的重要支流类群。同时发现中下游物种多样性高,上游及河口江段多样性低的格局。采用冗余分析方法(RDA)分析了鱼类多样性与环境因子的关系,发现年均气温、降雨量、年均径流量、河流宽度与透明度是珠江水系河流鱼类群落结构差异的主要影响因子,其中年均气温是影响鱼类群落分布的最关键因子之一。与历史资料对比后发现,珠江鱼类种类明显减少、空间分布也发生了巨大改变。研究是珠江水系野生渔业资源长期调查的一部分,研究结果将对渔业资源的多样性保护和可持续利用具有指导意义。
The Pearl River,which is 2400 km long and flows into the South China Sea,is the longest river in subtropical southern China. The warm humid climate of the region,which promotes high species diversity,has resulted in significant aquatic biodiversity and the development of lucrative commercial fisheries. Nevertheless,studies on the spatial distribution of fish communities in the Pearl River system are limited,especially,considering the decline in global fishery resources due to numerous anthropogenic stressors,such as overfishing,dam construction,introduction of invasive species,and climate change. Thus,studies on the spatial distribution of fish species in the Pearl River are urgently needed. This study investigated the spatial patterns of fish diversity and distribution in the Pearl River Basin based on an investigation of 13 sample sites. A total of 10,119 individual fishes were collected during the study period,comprising 94 taxa representing 17 families and 72 genera. The Cyprinidae family,representing 49 taxa,accounted for 52% of the total number of fish species collected. Cyprinidae were the dominant family in the Pearl River,followed by Bagridae( 7%; seven species) and Cobitidae( 4%; four species). Non-metric multidimensional scaling( NMDS) was applied to the collected data and three separate groups were identified. The spatial variation in upstream fish assemblages differed significantly from that in downstream assemblages( P < 0. 05),although clustering and diversity showed a significant spatial autocorrelation. A significantly high species diversity was recorded in the Pearl River,but low species diversity existed in the up and downstream reaches. Redundancy analysis was used to describe the relationship between the diversity patterns of fish species and associated environmental factors. Temperature,precipitation,water discharge,river width,and water chemistry play important roles in species diversity,with the average annual temperature accounting for the largest proportion of variance. A comparison with historical data for the Pearl River revealed a dramatic decline in fish species and a significant change in the spatial distribution of fish communities. Currently,the most serious threat to the ecology of the Pearl River network is the invasion of alien species. For example,non-native tilapia are now widely distributed in most of the Pearl River system,even surviving in the cooler,upper reaches of the Nanpanjiang River. In the Hongshuihe,Beijiang,and Dongjiang Rivers,tilapia is the dominant species recorded and has frequently been collected in the middle and lower reaches of the Pearl River. The present study is part of a long-term investigation into the wild fishery resources of the Pearl River. Therefore,understanding the processes outlined in this study will assist in the conservation of fish community diversity,which is critical to the success and sustainability of commercial fisheries in the Pearl River. Regularized discriminant analysis( RDA) is used by ecologists to relate a dataset( Y) of response variables( such as species abundance) to a second dataset( X) of explanatory variables( often environmental factors) and consistently outperformed other statistical tests using all the data in this study.
The Pearl River,which is 2400 km long and flows into the South China Sea,is the longest river in subtropical southern China. The warm humid climate of the region,which promotes high species diversity,has resulted in significant aquatic biodiversity and the development of lucrative commercial fisheries. Nevertheless,studies on the spatial distribution of fish communities in the Pearl River system are limited,especially,considering the decline in global fishery resources due to numerous anthropogenic stressors,such as overfishing,dam construction,introduction of invasive species,and climate change. Thus,studies on the spatial distribution of fish species in the Pearl River are urgently needed. This study investigated the spatial patterns of fish diversity and distribution in the Pearl River Basin based on an investigation of 13 sample sites. A total of 10,119 individual fishes were collected during the study period,comprising 94 taxa representing 17 families and 72 genera. The Cyprinidae family,representing 49 taxa,accounted for 52% of the total number of fish species collected. Cyprinidae were the dominant family in the Pearl River,followed by Bagridae( 7%; seven species) and Cobitidae( 4%; four species). Non-metric multidimensional scaling( NMDS) was applied to the collected data and three separate groups were identified. The spatial variation in upstream fish assemblages differed significantly from that in downstream assemblages( P < 0. 05),although clustering and diversity showed a significant spatial autocorrelation. A significantly high species diversity was recorded in the Pearl River,but low species diversity existed in the up and downstream reaches. Redundancy analysis was used to describe the relationship between the diversity patterns of fish species and associated environmental factors. Temperature,precipitation,water discharge,river width,and water chemistry play important roles in species diversity,with the average annual temperature accounting for the largest proportion of variance. A comparison with historical data for the Pearl River revealed a dramatic decline in fish species and a significant change in the spatial distribution of fish communities. Currently,the most serious threat to the ecology of the Pearl River network is the invasion of alien species. For example,non-native tilapia are now widely distributed in most of the Pearl River system,even surviving in the cooler,upper reaches of the Nanpanjiang River. In the Hongshuihe,Beijiang,and Dongjiang Rivers,tilapia is the dominant species recorded and has frequently been collected in the middle and lower reaches of the Pearl River. The present study is part of a long-term investigation into the wild fishery resources of the Pearl River. Therefore,understanding the processes outlined in this study will assist in the conservation of fish community diversity,which is critical to the success and sustainability of commercial fisheries in the Pearl River. Regularized discriminant analysis( RDA) is used by ecologists to relate a dataset( Y) of response variables( such as species abundance) to a second dataset( X) of explanatory variables( often environmental factors) and consistently outperformed other statistical tests using all the data in this study.
引文
[1]王忠锁,陈明华,吕偲,许崇任,雷光春.鄱阳湖银鱼多样性及其时空格局.生态学报,2006,26(5):1337-1344.
[2]Hanski I.Metapopulation Ecology.New York:Oxford University Press,1999.
[3]江洪,马克平,张艳丽,朱春全,Strittholt J R.基于空间分析的保护生物学研究.植物生态学报,2004,28(4):562-578.
[4]费骥慧,汪兴中,邵晓阳.洱海鱼类群落的空间分布格局.水产学报,2012,36(8):1225-1233.
[5]邓凯,李丽,吴巩胜,周跃,赵如.景观空间格局对滇金丝猴猴群分布的影响.生态学报,2014,34(17):4999-5006.
[6]刘吉平,吕宪国.三江平原湿地鸟类丰富度的空间格局及热点地区保护.生态学报,2011,31(20):5894-5902.
[7]Nakagawa H.Contribution of environmental and spatial factors to the structure of stream fish assemblages at different spatial scales.Ecology of Freshwater Fish,2014,23(2):208-223.
[8]Jackson D A,Peres-Neto P R,Olden J D.What controls who is where in freshwater fish communities-the roles of biotic,abiotic,and spatial factors.Canadian Journal of Fisheries and Aquatic Sciences,2001,58(1):157-170.
[9]Ostrand K G,Wilde G R.Seasonal and spatial variation in a prairie stream-fish assemblage.Ecology of Freshwater Fish,2002,11(3):137-149.
[10]Mason N W H,Lanoiselée C,Mouillot D,Irz P,Argillier C.Functional characters combined with null models reveal inconsistency in mechanisms of species turnover in lacustrine fish communities.Oecologia,2007,153(2):441-452.
[11]Mouillot D,Dumay O,Tomasini J A.Limiting similarity,niche filtering and functional diversity in coastal lagoon fish communities.Estuarine,Coastal and Shelf Science,2007,71(3/4):443-456.
[12]Sharma S,Legendre P,De Cáceres M,Boisclair D.The role of environmental and spatial processes in structuring native and non-native fish communities across thousands of lakes.Ecography,2011,34(5):762-771.
[13]Grenouillet G,Pont D,HérisséC.Within-basin fish assemblage structure:the relative influence of habitat versus stream spatial position on local species richness.Canadian Journal of Fisheries and Aquatic Sciences,2004,61(1):93-102.
[14]Peres-Neto P,Legendre P,Dray S,Borcard D.Variation partitioning of species data matrices:estimation and comparison of fractions.Ecology,2006,87(10):2614-2625.
[15]Poff N L,Allan J D.Functional organization of stream fish assemblages in relation to hydrological variability.Ecology,1995,76(2):606-627.
[16]Sala O E,Chapin III F S,Armesto J J,Berlow E,Bloomfield J,Dirzo R,Huber-Sanwald E,Huenneke L F,Jackson R B,Kinzig A,Leemans R,Lodge D M,Mooney H A,Oesterheld M,Poff N L,Sykes M T,Walker B H,Walker M,Wall D H.Global biodiversity scenarios for the year2100.Science,2000,287(5459):1770-1774.
[17]Dulvy N K,Sadovy Y,Reynolds J D.Extinction vulnerability in marine populations.Fish and Fisheries,2003,4(1):25-64.
[18]Tolimieri N,Levin P S.Assemblage structure of Eastern Pacific groundfishes on the U.S.continental slope in relation to physical and environmental variables.Transactions of the American Fisheries Society,2006,135(2):317-332.
[19]李圣法,程家骅,严利平.东海大陆架鱼类群落的空间结构.生态学报,2007,27(11):4377-4386.
[20]傅萃长.长江流域鱼类多样性空间格局与资源分析——兼论银鱼的生物多样性与系统发育[D].上海:复旦大学,2003.
[21]苏巍.海州湾海域鱼类群落多样性及其与环境因子的关系[D].青岛:中国海洋大学,2014.
[22]张殷波,郭柳琳,王伟,田瑜,李俊生.秦岭重点保护植物丰富度空间格局与热点地区.生态学报,2014,34(8):2109-2117.
[23]陆奎贤.珠江水系渔业资源.广州:广东科技出版社,1990.
[24]Horton R E.Erosional development of streams and their drainage basins;hydrophysical approach to quantitative morphology.Geological Society of America Bulletin,1945,56(3):275-370.
[25]潘炯华,钟麟,郑慈英,伍汉霖,刘家照.广东淡水鱼类志.广州:广东科技出版社,1991.
[26]广西壮族自治区水产研究所,中国科学院动物研究所.广西淡水鱼类志(第二版).南宁:广西人民出版社,2006.
[27]郑慈英.珠江鱼类志.北京:科技出版社,1989.
[28]Legendre P,Legendre L.Numerical Ecology.3rd ed.Amsterdam,the Netherlands:Elsevier Science,2012.
[29]Walters D M,Leigh D S,Freeman M C,Freeman B J,Pringle C M.Geomorphology and fish assemblages in a Piedmont river basin,U.S.A.Freshwater Biology,2003,48(11):1950-1970.
[30]Kilroy C,Larned S T,Biggs B J F.The non-indigenous diatom Didymosphenia geminata alters benthic communities in New Zealand rivers.Freshwater Biology,2009,54(9):1990—2002.
[31]Makarenkov V,Legendre P.Nonlinear redundancy analysis and canonical correspondence analysis based on polynomial regression.Ecology,2002,83(4):1146-1161.
[32]Legendre P,Gallagher E D.Ecologically meaningful transformations for ordination of species data.Oecologia,2001,129(2):271-280.
[33]马克平,刘玉明.生物群落多样性的测定方法Ⅰ:α多样性的测度方法(下).生物多样性,1994,2(4):231-239.
[34]谭细畅,李新辉,李跃飞,李捷.尼罗罗非鱼早期发育形态及其在珠江水系的空间分布.生物安全学报,2012,21(4):295-299.
[35]顾党恩,牟希东,罗渡,李莹莹,杨叶欣,徐猛,胡隐昌.广东省主要水系罗非鱼的建群状况.生物安全学报,2012,21(4):277-282.
[36]Shuai F M,Li X H,Li Y F,Li J,Yang J P,Lek S.Forecasting the invasive potential of Nile tilapia(Oreochromis niloticus)in a large subtropical river using a univariate approach.Fundamental and Applied Limnology,2015,187(2):165-176.
[37]Oberdorff T,Hugueny B,Vigneron T.Is assemblage varia bility related to environmental variability?An answer for riverine fish.Oikos,2001,93(3):419-428.
[38]Pyron M,Lauer T E.Hydrological variation and fish assemblage structure in the middle Wabash River.Hydrobiologia,2004,525(1/3):203-213.
[39]Kouamélan E P,Teugels G G,N'Douba V,GooréBi G,KonéT.Fish diversity and its relationships with environmental variables in a West African basin.Hydrobiologia,2003,505(1/3):139-146.
[40]Brown L R.Fish communities and their associations with environmental variables,lower San Joaquin River drainage,California.Environmental Biology of Fishes,2000,57(3):251-269.
[41]Kadye W T,Magadza C H D,Moyo N A G,Kativu S.Stream fish assemblages in relation to environmental factors on a montane plateau(Nyika Plateau,Malawi).Environmental Biology of Fishes,2008,83(4):417-428.
[42]Ornellas A B,Coutinho R.Spatial and temporal patterns of distribution and abundance of a tropical fish assemblage in a seasonal Sargassum bed,Cabo Frio Island,Brazil.Journal of Fish Biology,1998,53:198-208.
[43]Tan X C,Li X H,Lek S,Li Y F,Wang C,Li J,Luo J R.Annual dynamics of the abundance of fish larvae and its relationship with hydrological variation in the Pearl River.Environmental Biology of Fishes,2010,88(3):217-225.
[44]王志恒,唐志尧,方精云.物种多样性地理格局的能量假说.生物多样性,2009,17(6):613-624.
[45]Allen A P,Gillooly J F.Assessing latitudinal gradients in speciation rates and biodiversity at the global scale.Ecology Letter,2006,9(8):947-954.
[46]Currie D J,Francis A P.Regional versus climatic effect on taxon richness in angiosperms:reply to Qian and Ricklefs.The American Naturalist,2004,163(5):780-785.
[2]Hanski I.Metapopulation Ecology.New York:Oxford University Press,1999.
[3]江洪,马克平,张艳丽,朱春全,Strittholt J R.基于空间分析的保护生物学研究.植物生态学报,2004,28(4):562-578.
[4]费骥慧,汪兴中,邵晓阳.洱海鱼类群落的空间分布格局.水产学报,2012,36(8):1225-1233.
[5]邓凯,李丽,吴巩胜,周跃,赵如.景观空间格局对滇金丝猴猴群分布的影响.生态学报,2014,34(17):4999-5006.
[6]刘吉平,吕宪国.三江平原湿地鸟类丰富度的空间格局及热点地区保护.生态学报,2011,31(20):5894-5902.
[7]Nakagawa H.Contribution of environmental and spatial factors to the structure of stream fish assemblages at different spatial scales.Ecology of Freshwater Fish,2014,23(2):208-223.
[8]Jackson D A,Peres-Neto P R,Olden J D.What controls who is where in freshwater fish communities-the roles of biotic,abiotic,and spatial factors.Canadian Journal of Fisheries and Aquatic Sciences,2001,58(1):157-170.
[9]Ostrand K G,Wilde G R.Seasonal and spatial variation in a prairie stream-fish assemblage.Ecology of Freshwater Fish,2002,11(3):137-149.
[10]Mason N W H,Lanoiselée C,Mouillot D,Irz P,Argillier C.Functional characters combined with null models reveal inconsistency in mechanisms of species turnover in lacustrine fish communities.Oecologia,2007,153(2):441-452.
[11]Mouillot D,Dumay O,Tomasini J A.Limiting similarity,niche filtering and functional diversity in coastal lagoon fish communities.Estuarine,Coastal and Shelf Science,2007,71(3/4):443-456.
[12]Sharma S,Legendre P,De Cáceres M,Boisclair D.The role of environmental and spatial processes in structuring native and non-native fish communities across thousands of lakes.Ecography,2011,34(5):762-771.
[13]Grenouillet G,Pont D,HérisséC.Within-basin fish assemblage structure:the relative influence of habitat versus stream spatial position on local species richness.Canadian Journal of Fisheries and Aquatic Sciences,2004,61(1):93-102.
[14]Peres-Neto P,Legendre P,Dray S,Borcard D.Variation partitioning of species data matrices:estimation and comparison of fractions.Ecology,2006,87(10):2614-2625.
[15]Poff N L,Allan J D.Functional organization of stream fish assemblages in relation to hydrological variability.Ecology,1995,76(2):606-627.
[16]Sala O E,Chapin III F S,Armesto J J,Berlow E,Bloomfield J,Dirzo R,Huber-Sanwald E,Huenneke L F,Jackson R B,Kinzig A,Leemans R,Lodge D M,Mooney H A,Oesterheld M,Poff N L,Sykes M T,Walker B H,Walker M,Wall D H.Global biodiversity scenarios for the year2100.Science,2000,287(5459):1770-1774.
[17]Dulvy N K,Sadovy Y,Reynolds J D.Extinction vulnerability in marine populations.Fish and Fisheries,2003,4(1):25-64.
[18]Tolimieri N,Levin P S.Assemblage structure of Eastern Pacific groundfishes on the U.S.continental slope in relation to physical and environmental variables.Transactions of the American Fisheries Society,2006,135(2):317-332.
[19]李圣法,程家骅,严利平.东海大陆架鱼类群落的空间结构.生态学报,2007,27(11):4377-4386.
[20]傅萃长.长江流域鱼类多样性空间格局与资源分析——兼论银鱼的生物多样性与系统发育[D].上海:复旦大学,2003.
[21]苏巍.海州湾海域鱼类群落多样性及其与环境因子的关系[D].青岛:中国海洋大学,2014.
[22]张殷波,郭柳琳,王伟,田瑜,李俊生.秦岭重点保护植物丰富度空间格局与热点地区.生态学报,2014,34(8):2109-2117.
[23]陆奎贤.珠江水系渔业资源.广州:广东科技出版社,1990.
[24]Horton R E.Erosional development of streams and their drainage basins;hydrophysical approach to quantitative morphology.Geological Society of America Bulletin,1945,56(3):275-370.
[25]潘炯华,钟麟,郑慈英,伍汉霖,刘家照.广东淡水鱼类志.广州:广东科技出版社,1991.
[26]广西壮族自治区水产研究所,中国科学院动物研究所.广西淡水鱼类志(第二版).南宁:广西人民出版社,2006.
[27]郑慈英.珠江鱼类志.北京:科技出版社,1989.
[28]Legendre P,Legendre L.Numerical Ecology.3rd ed.Amsterdam,the Netherlands:Elsevier Science,2012.
[29]Walters D M,Leigh D S,Freeman M C,Freeman B J,Pringle C M.Geomorphology and fish assemblages in a Piedmont river basin,U.S.A.Freshwater Biology,2003,48(11):1950-1970.
[30]Kilroy C,Larned S T,Biggs B J F.The non-indigenous diatom Didymosphenia geminata alters benthic communities in New Zealand rivers.Freshwater Biology,2009,54(9):1990—2002.
[31]Makarenkov V,Legendre P.Nonlinear redundancy analysis and canonical correspondence analysis based on polynomial regression.Ecology,2002,83(4):1146-1161.
[32]Legendre P,Gallagher E D.Ecologically meaningful transformations for ordination of species data.Oecologia,2001,129(2):271-280.
[33]马克平,刘玉明.生物群落多样性的测定方法Ⅰ:α多样性的测度方法(下).生物多样性,1994,2(4):231-239.
[34]谭细畅,李新辉,李跃飞,李捷.尼罗罗非鱼早期发育形态及其在珠江水系的空间分布.生物安全学报,2012,21(4):295-299.
[35]顾党恩,牟希东,罗渡,李莹莹,杨叶欣,徐猛,胡隐昌.广东省主要水系罗非鱼的建群状况.生物安全学报,2012,21(4):277-282.
[36]Shuai F M,Li X H,Li Y F,Li J,Yang J P,Lek S.Forecasting the invasive potential of Nile tilapia(Oreochromis niloticus)in a large subtropical river using a univariate approach.Fundamental and Applied Limnology,2015,187(2):165-176.
[37]Oberdorff T,Hugueny B,Vigneron T.Is assemblage varia bility related to environmental variability?An answer for riverine fish.Oikos,2001,93(3):419-428.
[38]Pyron M,Lauer T E.Hydrological variation and fish assemblage structure in the middle Wabash River.Hydrobiologia,2004,525(1/3):203-213.
[39]Kouamélan E P,Teugels G G,N'Douba V,GooréBi G,KonéT.Fish diversity and its relationships with environmental variables in a West African basin.Hydrobiologia,2003,505(1/3):139-146.
[40]Brown L R.Fish communities and their associations with environmental variables,lower San Joaquin River drainage,California.Environmental Biology of Fishes,2000,57(3):251-269.
[41]Kadye W T,Magadza C H D,Moyo N A G,Kativu S.Stream fish assemblages in relation to environmental factors on a montane plateau(Nyika Plateau,Malawi).Environmental Biology of Fishes,2008,83(4):417-428.
[42]Ornellas A B,Coutinho R.Spatial and temporal patterns of distribution and abundance of a tropical fish assemblage in a seasonal Sargassum bed,Cabo Frio Island,Brazil.Journal of Fish Biology,1998,53:198-208.
[43]Tan X C,Li X H,Lek S,Li Y F,Wang C,Li J,Luo J R.Annual dynamics of the abundance of fish larvae and its relationship with hydrological variation in the Pearl River.Environmental Biology of Fishes,2010,88(3):217-225.
[44]王志恒,唐志尧,方精云.物种多样性地理格局的能量假说.生物多样性,2009,17(6):613-624.
[45]Allen A P,Gillooly J F.Assessing latitudinal gradients in speciation rates and biodiversity at the global scale.Ecology Letter,2006,9(8):947-954.
[46]Currie D J,Francis A P.Regional versus climatic effect on taxon richness in angiosperms:reply to Qian and Ricklefs.The American Naturalist,2004,163(5):780-785.