华南沿海日本大叶藻种群遗传结构和基因流格局
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摘要
海草床是全球三大海洋生态系统之一,具有重要的生态系统服务功能。然而受人为干扰和自然环境恶化的影响,海草床正面临全球性的衰退,因此对于海草床的保护和恢复刻不容缓。日本大叶藻是亚洲温带和热带地区广泛分布的海草优势种,在海草床生态系统服务功能中发挥着重要作用,本论文开发并筛选出适用的特异性微卫星引物,对华南沿海的日本大叶藻种群遗传结构进行研究,阐明克隆性和基因流在海草遗传多样性维持中的影响。主要研究结果及结论如下：
1.利用生物素链霉亲和素捕获法在日本大叶藻中筛选出12对多态微卫星引物,且PCR扩增重复性和稳定性很好,能够用于该物种种群遗传学研究。这12个位点在海南2个日本大叶藻种群57个样品中分别有2～10个等位基因,2个种群的平均等位基因数分别为4.1(BG)和4.4(DJ),观察和期望杂合度的范围分别为0.0000～1.0000、0.0000～0.7983(BG)和0.0000～1.00、0.0000～0.8542(DJ)。
2.在12个位点中,有2个位点(ZJ48和ZJ1368)受分化选择作用。利用其余10个中性位点对华南沿海8个日本大叶藻种群251个样品进行分析,结果表明8个种群中克隆丰富度最小为YT种群(0.742),最大为SX种群(0.980)；期望杂合度为0.373(YT)～0.621(DJ),显示出高的遗传多样性。种群间标准化的遗传分化系数F'ST为0.354。Structure、PCoA和AMOVA分析结果显示8个种群可聚成2个类群：广西的4个种群(SX、DC、GS和JD)聚为西部类群,海南(BG和DJ)和广东(YT和SZ)4个种群聚为东部类群,同时发现种群间存在距离隔离格局(IBD)。此外,类群内的基因流大于类群间,并且西部类群内的基因流大于东部类群。而广东的YT和SZ种群间存在遗传间断。
3.日本大叶藻种群克隆丰富度较高,种群内不存在自交情况,克隆子界为5.1m。通过对日本大叶藻种群10m×10m空间自相关分析表明,分株和中心基株水平在第一距离级内(0～1m)均存在显著的空间遗传结构(p<0.001),并且分株水平的空间遗传结构显著强于基株水平(co=17.373,p=0.023)；但随着距离的增加,分株和基株水平空间遗传结构之间的差异逐渐减小。这表明克隆性在较短距离内对空间遗传结果存在显著的影响。另外,有性繁殖的扩散方差(δ=3.18)远大于克隆扩散方差(δ=0.18),表明克隆生长在日本大叶藻种群内受到限制,较长距离的扩散仍依赖于有性繁殖。
As one of the three main ocean ecosystems, seagrass meadows have important ecosystem services. However, nowadays they are suffering from globally declining due to natural environment deterioration and human disturbance. Zostera japonica, a monoecious seagrass, is a common dominated species of seagrass beds. In this study, I screened12polymorphic loci for Z. japonica. Population genetic structure and genetic diversity of this species were clarified based on these loci. Furthermore, we tried to discern the roles of clonality and gene flow on genetic diversity in Z. japonica. The follwoings are three main conclusions:
1. Using the method of biotin-streptavidin, we obtained12polymorphic micro satellite loci in Z. japonica. These loci had excellent repeatability and stability in PCR, providing a powerful tool to study population genetics for the species.2-10alleles per loci were found in57samples of two Z. japonica populations (BG and DJ) located in Hainan province. The number of mean alleles in BG and DJ were4.1and4.4, respectively. Ranges of observed and expected heterozygosities were0.0000-1.0000,0.0000-0.7983(BG) and0.0000-1.00,0.0000-0.8542(DJ).
2. Among the12microsatellites loci,2loci (ZJ48and ZJ1368) were not neutral.251samples collected from8Z. japonica populations were genotyped using10neutral loci. The minimum clonal richness was0.742(YT) and the maximum was0.980(SZ); the expected heterozygosity ranged from0.373(YT) to0.621(DJ). The standard genetic differentiation coefficient (FSr) was0.354among all populations. Using Structure, PCoA and AMOVA, the8populations could be clustered into2groups:the western group (SX, DC, GS and JD) and the eastern group (BG, DJ, YT and SZ). There was a significant relationship between the shortest ocean surface distance and genetic distance, indicating an isolation-by-distance pattern. Gene flow within western group was larger than that of the eastern one, while gene flows within groups were larger than between groups. A genetic barrier was identified between populations YT and SZ.
3. Clonal richness was high in Z. japonica, and selfing was absent within population. The clonal sub-range was5.1m. A significant difference in spatial genetic structure at the first distance class (0-1m) was observed between ramet-level and genet-level. The spatial genetic structure of ramet-level was much stronger than that of genet-level (ω=17.373, p=0.023). As distance increasing, difference in spatial genetic structure between the two levels decreased gradually. Dispersal variance of sexual reproduction (σ=3.18) was larger than that of clonal dispersal (σ=0.18), suggesting that clone growth was restricted and dispersal of long distance depended on sexual reproduction.
1.利用生物素链霉亲和素捕获法在日本大叶藻中筛选出12对多态微卫星引物,且PCR扩增重复性和稳定性很好,能够用于该物种种群遗传学研究。这12个位点在海南2个日本大叶藻种群57个样品中分别有2～10个等位基因,2个种群的平均等位基因数分别为4.1(BG)和4.4(DJ),观察和期望杂合度的范围分别为0.0000～1.0000、0.0000～0.7983(BG)和0.0000～1.00、0.0000～0.8542(DJ)。
2.在12个位点中,有2个位点(ZJ48和ZJ1368)受分化选择作用。利用其余10个中性位点对华南沿海8个日本大叶藻种群251个样品进行分析,结果表明8个种群中克隆丰富度最小为YT种群(0.742),最大为SX种群(0.980)；期望杂合度为0.373(YT)～0.621(DJ),显示出高的遗传多样性。种群间标准化的遗传分化系数F'ST为0.354。Structure、PCoA和AMOVA分析结果显示8个种群可聚成2个类群：广西的4个种群(SX、DC、GS和JD)聚为西部类群,海南(BG和DJ)和广东(YT和SZ)4个种群聚为东部类群,同时发现种群间存在距离隔离格局(IBD)。此外,类群内的基因流大于类群间,并且西部类群内的基因流大于东部类群。而广东的YT和SZ种群间存在遗传间断。
3.日本大叶藻种群克隆丰富度较高,种群内不存在自交情况,克隆子界为5.1m。通过对日本大叶藻种群10m×10m空间自相关分析表明,分株和中心基株水平在第一距离级内(0～1m)均存在显著的空间遗传结构(p<0.001),并且分株水平的空间遗传结构显著强于基株水平(co=17.373,p=0.023)；但随着距离的增加,分株和基株水平空间遗传结构之间的差异逐渐减小。这表明克隆性在较短距离内对空间遗传结果存在显著的影响。另外,有性繁殖的扩散方差(δ=3.18)远大于克隆扩散方差(δ=0.18),表明克隆生长在日本大叶藻种群内受到限制,较长距离的扩散仍依赖于有性繁殖。
As one of the three main ocean ecosystems, seagrass meadows have important ecosystem services. However, nowadays they are suffering from globally declining due to natural environment deterioration and human disturbance. Zostera japonica, a monoecious seagrass, is a common dominated species of seagrass beds. In this study, I screened12polymorphic loci for Z. japonica. Population genetic structure and genetic diversity of this species were clarified based on these loci. Furthermore, we tried to discern the roles of clonality and gene flow on genetic diversity in Z. japonica. The follwoings are three main conclusions:
1. Using the method of biotin-streptavidin, we obtained12polymorphic micro satellite loci in Z. japonica. These loci had excellent repeatability and stability in PCR, providing a powerful tool to study population genetics for the species.2-10alleles per loci were found in57samples of two Z. japonica populations (BG and DJ) located in Hainan province. The number of mean alleles in BG and DJ were4.1and4.4, respectively. Ranges of observed and expected heterozygosities were0.0000-1.0000,0.0000-0.7983(BG) and0.0000-1.00,0.0000-0.8542(DJ).
2. Among the12microsatellites loci,2loci (ZJ48and ZJ1368) were not neutral.251samples collected from8Z. japonica populations were genotyped using10neutral loci. The minimum clonal richness was0.742(YT) and the maximum was0.980(SZ); the expected heterozygosity ranged from0.373(YT) to0.621(DJ). The standard genetic differentiation coefficient (FSr) was0.354among all populations. Using Structure, PCoA and AMOVA, the8populations could be clustered into2groups:the western group (SX, DC, GS and JD) and the eastern group (BG, DJ, YT and SZ). There was a significant relationship between the shortest ocean surface distance and genetic distance, indicating an isolation-by-distance pattern. Gene flow within western group was larger than that of the eastern one, while gene flows within groups were larger than between groups. A genetic barrier was identified between populations YT and SZ.
3. Clonal richness was high in Z. japonica, and selfing was absent within population. The clonal sub-range was5.1m. A significant difference in spatial genetic structure at the first distance class (0-1m) was observed between ramet-level and genet-level. The spatial genetic structure of ramet-level was much stronger than that of genet-level (ω=17.373, p=0.023). As distance increasing, difference in spatial genetic structure between the two levels decreased gradually. Dispersal variance of sexual reproduction (σ=3.18) was larger than that of clonal dispersal (σ=0.18), suggesting that clone growth was restricted and dispersal of long distance depended on sexual reproduction.
引文
Abe M, Yokota K, Kurashima A, Maegawa M (2009a) High water temperature tolerance in photosynthetic activity of Zostera japonica Ascherson & Graebner seedlings from Ago Bay, Mie Prefecture, central Japan. Fisheries Science 75,1117-1123.
Abe M, Yokota K, Kurashima A, Maegawa M (2009b) Temperature characteristics in seed germination and growth of Zostera japonica Ascherson & Graebner from Ago Bay, Mie Prefecture, central Japan. Fisheries Science 75,921-927.
Aioi K, Nakaoka M (2003) The Seagrass of Japan. In:Green EP and Short FT (eds) World Atlas of Seagrasses. University of California Press, Berkeley.
Alberto F, Correia L, Arnaud-Haond S, Billot C, Duarte CM, Serrao E (2003a) New microsatellite markers for the endemic Mediterranean seagrass Posidonia oceanica. Molecular Ecology Notes 3,253-255.
Alberto F, Correia L, Billot C, Duarte CM, Serrao E (2003b) Isolation and characterization of microsatellite markers for the seagrass Cymodocea nodosa. Molecular Ecology Notes 3, 397-399.
Alberto F, Gouveia L, Arnaud-Haond S, Perez-Llorens JL, Duarte CM, Serrao EA (2005) Within-population spatial genetic structure, neighbourhood size and clonal subrange in the seagrass Cymodocea nodosa. Molecular Ecology 14,2669-2681.
Alberto F, Massa S, Manent P, Diaz-Almela E, Arnaud-Haond S, Duarte CM, Serrao EA (2008) Genetic differentiation and secondary contact zone in the seagrass Cymodocea nodosa across the Mediterranean-Atlantic transition regions. Journal of Biogeography 35, 1279-1294.
Allendorf F W, Luikart G (2007) Conservation and the Genetics of Populations. Blackwell.
Anderson JB, Kohn LM (1995) Clonality in soilborne, plant-pathogenic fungi. Annual Review of Phytopathology 33,369-391.
Angevine MW, Handel SN (1986) Invasion of forest floor space, clonal architecture, and population growth in the perennial herb Clintonia Borealis. Journal of Ecology 74, 547-560.
Araki S, Kunii H (2006) Allozymic implications of the propagation of eelgrass Zostera japonica within a river system. Limnology 7,15-21.
Arnaud-Haond S, Alberto F, Teixeira S, Procaccini G, Serrao EA, Duarte CM (2005) Assessing genetic diversity in clonal organisms:low diversity or low resolution? Combining power and cost efficiency in selecting markers. Journal of Heredity 96,434-440.
Arnaud-Haond S, Belkhir K (2007) GENCLONE:a computer program to analyse genotypic data, test for clonality and describe spatial clonal organization. Molecular Ecology Notes 7, 15-17.
Arnaud-Haond S, Duarte CM, Alberto F, Serrao EA (2007a) Standardizing methods to address clonality in population studies. Molecular Ecology 16,5115-5139.
Arnaud-Haond S, Duarte CM, Diaz-Aimela E, Marba N, Sintes T, Sen-ao EA(2012) Implications of extreme life span in clonal organisms:millenary clones in meadows of the threatened seagrass Posidonia oceanica. PLoS ONE 7, e30454.
Arnaud-Haond S, Migliaccio M, Diaz-Almela E, Teixeira S, van de Vliet MS, Alberto F, Procaccini G, Duarte CM, Serrao EA (2007b) Vicariance patterns in the Mediterranean Sea:east-west cleavage and low dispersal in the endemic seagrass Posidonia oceanica. Journal of Biogeography 34,963-976.
Avise JC (2004) Molecular Markers, Natural History and Evolution. Sinauer Associates, Sunderland, Mass.
Ballesteros E, Cebrian E, Garcia-Rubies A, Alcoverro T, Romero J, Font X (2005) Pseudovivipary, a new fonn of asexual reproduction in the seagrass Posidonia oceanica. Botanica Marina 48,175-177.
Beaumont MA, Nichols RA(1996) Evaluating loci for use in the genetic analysis of population structure. Proceedings of the Royal Society of London Series B:Biological Sciences 263, 1619-1626.
Beebee T, Rowe G (2004) An Introduction to Molecular Ecology. Oxford University Press.
Beerli R (2011) Migrate-N, Version 3.2.17. http://popgen.scs.fsu.edu/mirate-n.html.
Bell AD, Tomlinson PB (1980) Adaptive architecture in rhizomatous plants. Botanical Journal of the Linnean Society 80,125-160.
Benjamin KJ, Walker DI, McComb AJ, Kuo J (1999) Structural response of marine and estuarine plants of Halophila ovalis (R-Br.) Hook. f. to long-term hyposalinity. Aquatic Botany 64, 1-17.
Billingham MR, Simoes T, Reusch TBH, Serrao EA(2007) Genetic sub-structure and intermediate optimal outcrossing distance in the marine angiosperm Zostera marina. Marine Biology 152,793-801.
Bohonak AJ (2002) IBD (Isolation by Distance):a program for analyses of isolation by distance. Journal of Heredity 93,153-154.
Brun FG, Cummaudo F, Olive I,Vergara JJ, Perez-Llorens JL (2007) Clonal extent, apical dominance and networking features in the phalanx angiosperm Zostera noltii Hornem. Marine Biology 151,1917-1927.
Cain ML, Dudle DA, Evans JP (1996) Spatial models of foraging in clonal plant species. American Journal of Botany 83,76-85.
Campbell SJ, McKenzie LJ, Kerville SP (2006) Photosynthetic responses of seven tropical seagrasses to elevated seawater temperature. Journal of Experimental Marine Biology and Ecology 330,455-468.
Capiomont A, Sandmeier M, Caye Q Meinesz A (1996) Enzyme polymorphism in Posidonia oceanica, a seagrass endemic to the Mediterranean. Aquatic Botany 54,265-277.
Casa AM, Mitchell SE, Hamblin MT, Sun H, Bowers JE, Paterson AH, Aquadro CF, Kresovich S (2005) Diversity and selection in sorghum:simultaneous analyses using simple sequence repeats. Theoretical and Applied Genetics 111,23-30.
Ceccherelli G, Campo D, Milazzo M (2007) Short-term response of the slow growing seagrass Posidonia oceanica to simulated anchor impact. Marine Environmental Research 63, 341-349.
Cornuet JM, Luikart G (1996) Description and power analysis of two tests for detecting recent population bottlenecks from allele frequency data. Genetics 144,2001-2014.
Coyer JA, Miller KA, Engle JM, Veldsink J, Cabello-Pasini A, Stam WT, Olsen JL (2008) Eelgrass meadows in the California channel islands and adjacent coast reveal a mosaic of two species, evidence for introgression and variable clonality. Annals of Botany 101, 73-87.
Coyer JA, Reusch TBH, Stam WT, Serrao EA, Pearson Q Procaccinis Q Olsen JL (2004) Characterization of microsatellite loci in the dwarf eelgrass Zostera noltii (Zosteraceae) and cross-reactivity with Z. japonica. Molecular Ecology Notes 4,497-499.
Cristescu R, Sherwin WB, Handasyde K, Cahill V, Cooper DW (2010) Detecting bottlenecks using BOTTLENECK 1.2.02 in wild populations:the importance of the microsatellite structure. Conservation Genetics 11,1043-1049.
Davis MB, Shaw RG (2001) Range shifts and adaptive responses to Quaternary climate change. Science 292,673-679.
Di Rienzo A, Peterson AC, Garza JC,'Valdes AM, Slatkin M, Freimer NB (1994) Mutational processes of simple-sequence repeat loci in human populations. Proceedings of the National Academy of Sciences of the United States of America 91,3166-3170.
Diaz-Almela E, Arnaud-Haond S, Vliet MS, Alvarez E, Marba N, Duarte CM, Serrao EA(2007) Feed-backs between genetic structure and perturbation-driven decline in seagrass (Posidonia oceanica) meadows. Conservation Genetics 8,1377-1391.
Diekmann OE, Coyer JA, Ferreira J, Olsen JL, Stam WT, Pearson GA, Serrao EA (2005) Popoulation genetics of Zostera noltii along the west Iberian coast:consequences of small population size, habitat discontinuity and near-shore currents. Marine Ecology Progress Series 290,89-96.
Dorken ME, Eckert CG (2001) Severely reduced sexual reproduction in northern populations of a clonal plant, Decodon verticillatus (Lythraceae). Journal of Ecology 89,339-350.
Douhovnikoff V, Dodd RS (2003) Intra-clonal variation and a similarity threshold for identification of clones:application to Salix exigua using AFLP molecular markers. Theoretical and Applied Genetics 106,1307-1315.
Duarte CM (1991) Allometric scaling of seagrass form and productivity. Marine Ecology Progress Series 77,289-300.
Duarte CM, Marba N, Krause-Jensen D, Sanchez-Camacho M (2007) Testing the predictive power of seagrass depth limit models. Estuaries andCoasts 30,652-656.
Epperson BK (1993) Recent advances in correlation studies of spatial patterns of genetic-variation. Evolutionary Biology 27,95-155.
Epperson BK (2003) Geographical Genetics:Monographs in Population Biology. Princeton University Press, Princeton, New Jersey.
Erftemeijer PLA, Stapel J (1999) Primary production of deep-water Halophila ovalis meadows. Aquatic Botany 65,71-82.
Evanno G, Regnaut S, Goudet J (2005) Detecting the number of clusters of individuals using the software STRUCTURE:a simulation study. Molecular Ecology 14,2611-2620.
Fenster CB,Vekemans X, Hardy OJ (2003) Quantifying gene flow from spatial genetic structure data in a metapopulation of Chamaecrista fasciculata (Leguminosae). Evolution 57, 995-1007.
Flint J, Bond J, Rees DC, Boyce AJ, Roberts-Thomson JM, Excoffier L, Clegg JB, Beaumont MA, Nichols RA, Harding RM (1999) Minisatellite mutational processes reduce Fst estimates. Human Genetics 105,567-576.
Gao H, Jiang K, Geng Y, Chen XY (2012) Development of microsatellite primers of the largest seagrass, Enhalus acoroides (Hydrocharitaceae). American Journal of Botany 99, E99-E101.
Gliddon C, Belhassen E, Gouyou PH (1987) Genetic neighborhoods in plants with diverse systems of mating and different patterns of growth. Heredity 59.29-32.
Goudet J (2002) FSTAT (Version 2.9.3.2):A computer program to calculate F-statistics. Updated from Goudet (1995). Available from http://www.unil.ch/izea/softwares/fstat.html.Journal of Heredity 86,485-486.
Halkett F, Simon JC, Balloux F (2005) Tackling the population genetics of clonal and partially clonal organisms. Trends in Ecology & Evolution 20,194-201.
Halpern BS, Walbridge S, Selkoe KA, Kappel CV, Micheli F, D'Agrosa C, Bruno JF, Casey KS, Ebert C, Fox HE, Fujita R, Heinemann D, Lenihan HS, Mad in EMP, Perry MT, Selig ER, Spalding M, Steneck R, Watson R (2008) A global map of human impact on marine ecosystems. Science 319,948-952.
Hammerli A, Reusch TBH (2003a) Genetic neighbourhood of clone structures in eelgrass meadows quantified by spatial autocorrelation of microsatellite markers. Heredity 91, 448-455.
Hammerli A, Reusch TBH (2003b) Inbreeding depression influences genet size distribution in a marine angiosperm. Molecular Ecology 12,619-629.
Harada Y, Iwasa Y(1996) Analyses of spatial patterns and population processes of clonal plants. Researches on Population Ecology (Kyoto) 38,153-164.
Hardy OJ, Vekemans X (1999) Isolation by distance in a continuous population:reconciliation between spatial autocorrelation analysis and population genetics models. Heredity 83, 145-154.
Hardy OJ, Vekemans X (2002) SPAGEDi:a versatile computer program to analyse spatial genetic structure at the individual or population levels. Molecular Ecology Notes 2,618-620.
Harrison PG, Bigley RE (1982) The recent introduction of the seagrass Zostera japonica Aschers. and Graebn. to the Pacific Coast of North America. Canadian Journal of Fisheries and Aquatic Sciences 39,1642-1648.
Hartog Cd, Kuo J (2006) Taxonomy and Biogeography of Seagrasses. In: Larkum, A.D.W., Orth, R.J., Duarte, CM. (Eds.), Seagrass:Biology, Ecology and Consei-vation. Springer, The Netherland.
Hedrick PW (2001) Conservation genetics:where are we now? Trends in Ecology & Evolution 16, 629-636.
Hedrick PW (2005) A standardized genetic differentiation measure. Evolution 59,1633-1638.
Hellberg ME (1995) Relationships between inferred levels of gene flow and geographic distance in a philopatric coral, Balanophyllia elegans. Evolution 48,1829-1854.
Hughes AR, Stachowicz JJ (2004) Genetic diversity enhances the resistance of a seagrass ecosystem to disturbance. Proceedings of the National Academy of Sciences of the United States of America 101,8998-9002.
Jiang K, Gao H, Xu NN, Tsang EPK, Chen XY(2011a) A set of microsatellite primers for Zostera japonica (Zosteraceae). American Journal of Botany 98, E236-E238.
Jiang K, Shi YS, Zhang J, Xu NN (2011b) Microsatellite primers for vulnerable seagrass Halophila beccarii (Hydrocharitaceae). American Journal of Botany 98, E155-E157.
Kendrick. GA, Duarte. CM, Marba. N (2005) Clonality in seagrasses, emergent properties and seagrass landscapes. Marine Ecology Progress Series 290,291-296.
Kornelis van Dijk J, van Tussenbroek BI (2010) Clonal diversity and structure related to habitat of the marine angiosperm Thalassia testudinum along the Atlantic coast of Mexico. Aquatic Botany 92,63-69.
Lalitha S (2000) Primer Primier 5. Biotech Software and Internet Report 1,270-273.
Lazrek F, Roussel V, Ronfort J, Cardinet Q Chardon F, Aouani M, Huguet T (2009) The use of neutral and non-neutral SSRs to analyse the genetic structure of a Tunisian collection of Medicago truncatula lines and to reveal associations with eco-environmental variables. Genetica 135,391-402.
Lee SY (1997) Annual cycle of biomass of a threatened population of the intertidal seagrass Zostera japonica in Hong Kong. Marine Biology 129,183-193.
Les DH, Cleland MA, Waycott M (1997) Phylogenetic studies in Alismatidae, Ⅱ:Evolution of marine angiosperms (seagrasses) and hydrophily. Systematic Botany 22,443-463.
Lirman D, Cropper WP (2003) The influence of salinity on seagrass growth, survivorship, and distribution within Biscayne Bay, Florida:Field, experimental, and modeling studies. Estuaries 26,131-141.
Lowe A, Harris S, Ashton P (2004) Ecological Genetics:Design, Analysis, and Application. Blackwell.
Lucas C, Thangaradjou T, Papenbrock J (2012) Development of a DNA barcoding system for seagrasses:successful but not simple. PLoSONE 7, e29987.
Manni F, Guerard E, Heyer E (2004) Geographic patterns of (genetic, morphologic, linguistic) variation:How barriers can be detected by using Monmonier's algorithm. Human Biology 76,173-190.
Marba N, Duarte CM (1998) Rhizome elongation and seagrass clonal growth. Marine Ecology Progress Series 174,269-280.
Marba N, Hemminga MA, Duarte CM (2006) Resource trans location within seagrass clones: allometric scaling to plant size and productivity. Oecologia 150,362-372.
Marba N, Hemminga MA, Mateo MA, Duarte CM, Mass YEM, Terrados J, Gacia E (2002) Carbon and nitrogen translocation between seagrass ramets. Marine Ecology Progress Series 226,287-300.
Meirmans PG (2006) Using the AMOVA framework to estimate a standardized genetic differentiation measure. Evolution 60,2399-2402.
Miller MP (1997) Tools for Population Genetic Analyses (TFPGA). A Windows program for the analysis of allozyme and molecular population genetic data version 1.3, Department of Biological Sciences, Northern Arizona University, Flagstaff, AZ.
Muniz-Salazar R, Talbot SL, Sage GK, Ward DH, Cabello-Pasini A (2005) Population genetic structure of annual and perennial populations of Zostera marina L. along the Pacific coast of Baja California and the Gulf of California. Molecular Ecology 14,711-722.
Nei M (1978) Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89,583-590.
Nielsen EE, Hansen MM, Meldrup D (2006) Evidence of microsatellite hitch-hiking selection in Atlantic cod (Gadus morhua L.):implications for inferring population structure in nonmodel organisms. Molecular Ecology 15,3219-3229.
Ohsawa T, Ide Y (2008) Global patterns of genetic variation in plant species along vertical and horizontal gradients on mountains. Global Ecology and Biogeography 17,152-163.
Peakall ROD, Smouse PE (2006) genalex 6:genetic analysis in Excel. Population genetic software for teaching and research. Molecular Ecology Notes 6,288-295.
Posey MH (1988) Community changes associated with the spread of an introduced seagrass, Zostera japonica. Ecology 69,974-983.
Prichard JK, Wen X, Falush D (2009) Documentation for structure software: version 2.3. http://pritch.bsd.uchicago.edu/structure.html.
Procaccini G, Mazzella L (1998) Population genetic structure and gene flow in the seagrass Posidonia oceanica assessed using microsatellite analysis. Marine Ecology Progress Series 169,133-141.
Procaccini G, Olsen JL, Reusch TBH (2007) Contribution of genetics and genomics to seagrass biology and conservation. Journal of Experimental Marine Biology and Ecology 350, 234-259.
Procaccini G, Orsini L, Ruggiero MV, Scardi M (2001) Spatial patterns of genetic diversity in Posidonia oceanica, an endemic Mediterranean seagrass. Molecular Ecology 10, 1413-1421.
Procaccini G, Piazzi L (2001) Genetic polymorphism and transplantation success in the mediterranean seagrass Posidonia oceanica. Restoration Ecology 9,332-338.
Procaccini G, Ruggiero MV, Orsini L (2002) Genetic structure and distribution of microsatellite population genetic diversity in Posidonia oceanica in the Mediterranean basin. Bulletin of Marine Science 71,1291-1297.
Procaccini G, Waycott M(1998) Microsatellite loci identified in the seagrass Posidonia oceanica (L.) Delile. Journal of Heredity 89,562-568.
Rannala B, Mountain JL (1997) Detecting immigration by using multilocus genotypes. Proceedings of the National Academy of Sciences of the United States of America 94, 9197-9201.
Reusch TBH (2000a) Five microsatellite loci in eelgrass Zostera marina and a test of cross-species amplification in Z. noltii and Z.japonica. Molecular Ecology 9,371-373.
Reusch TBH (2000b) Pollination in the marine realm:microsatellites reveal high outcrossing rates and multiple paternity in eelgrass Zostera marina. Heredity 85,459-464.
Reusch TBH (2001) New markers-old questions:population genetics of seagrasses. Marine Ecology Progress Series 211,261-274.
Reusch TBH, Ehlers A, Hammerli A, Worm B (2005) Ecosystem recovery after climatic extremes enhanced by genotypic diversity. Proceedings of the National Academy of Sciences of the United States of America 102,2826-2831.
Reusch TBH, Stam WT, Olsen JL (1999) Size and estimated age of genets in eelgrass, Zostera marina, assessed with microsatellite markers. Marine Biology 133,519-525.
Reusch TBH, Stam WT, Olsen JL(2000) Amicrosatellite-based estimation of clonal diversity and population subdivision in Zostera marina, a marine flowering plant. Molecular Ecology 9, 127-140.
Richardson CA, Kennedy H, Duarte CM. Kennedy DP, Proud SV (1999) Age and growth of the fan mussel Pinna nobilis from south-east Spanish Mediterranean seagrass (Posidonia oceanicu) meadows. Marine Biology 133,205-212.
Rousset F (1997) Genetic differentiation and estimation of gene flow from F-statistics under isolation by distance. Genetics 145,1219-1228.
Ruggiero MV, Capone S, Pirozzi P, Reusch TBH, Procaccini G (2005a) Mating system and clonal architecture: a comparative study in two marine angiosperms. Evolutionary Ecology 19, 487-499.
Ruggiero MV, Reusch TBH, Procaccini G (2004) Polymorphic microsatellite loci for the marine angiosperm Cymodocea nodosa. Molecular Ecology Notes 4,512-514.
Ruggiero MV, Reusch TBH, Procaccini G (2005b) Local genetic structure in a clonal dioecious angiosperm. Molecular Ecology 14,957-967.
Ruggiero MV, Turk R, Procaccini G (2002) Genetic identity and homozygosity in North-Adriatic populations of Posidonia oceanica:an ancient, post-glacial clone? Conservation Genetics 3,71-74.
Schlotterer C (2004) The evolution of molecular markers-just a matter of fashion? Nature Reviews Genetics 5,63-69.
Serra IA, Innocenti AM, Di Maida G, Calvo S, Migliaccio M, Zambianchi E, Pizzigalli C, Arnaud-Haond S, Duarte CM, Serrao EA, Procaccini G (2010) Genetic structure in the Mediterranean seagrass Posidonia oceanica:disentangling past vicariance events from contemporary patterns of gene flow. Molecular Ecology 19,557-568.
Serra IA, Procaccini G, Intrieri MC, Migliaccio M, Mazzuca S, Innocenti AM (2007) Comparison of lSSR and SSR markers for analysis of genetic diversity in the seagrass Posidonia oceanica. Marine Ecology Progress Series 338,71-79.
Shi MM, Michalski SG, Chen XY, Durka W (2011) Isolation by elevation:genetic structure at neutral and putatively non-neutral loci in a dominant tree of subtropical forests, Castanopsis eyrei. PLoSONE 6, e21302.
Shin H, Choi HK (1998) Taxonomy and distribution of Zostera (Zosteraceae) in eastern Asia, with special reference to Korea. Aquatic Botany 60,49-66.
Short F, Carruthers T, Dennison W, Waycott M (2007) Global seagrass distribution and diversity:a bioregional model. Journal of Experimental Marine Biology and Ecology 350,3-20.
Short FT, Polidoro B, Livingstone SR, Carpenter K.E, Bandeira S, Bujang JS, Calumpong HP, Carruthers TJB, Coles RG, Dennison WC, Erftemeijer PLA, Fortes MD, Freeman AS, Jagtap TG, Kamal AHM, Kendrick GA, Judson Kenworthy W, La Nafie YA, Nasution IM, Orth RJ, Prathep A, Sanciangco JC, Tussenbroek Bv, Vergara SG, Waycott M, Zieman JC (2011) Extinction risk assessment of the world's seagrass species. Biological Conservation 144,1961-1971.
Sinclair EA, Anthony J, Coupland GT, Waycott M, Barrett MD, Barrett RL, Cambridge ML, Wallace MJ, Dixon KW, Krauss SL, Kendrick GA (2009) Characterisation of polymorphic microsatellite markers in the widespread Australian seagrass, Posidonia australis Hook. f. (Posidoniaceae), with cross-amplification in the sympatric P. sinuosa. Conservation Genetics Resources 1,273-276.
Smouse PE, Peakall ROD, Gonzales EVA (2008) A heterogeneity test for fine-scale genetic structure. Molecular Ecology 17,3389-3400.
Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011) MEGA5:Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28,2731-2739.
Tanaka N, Demise T, Ishii M, Shoji Y, Nakaoka M (2011) Genetic structure and gene flow of eelgrass Zostera marina populations in Tokyo Bay, Japan:implications for their restoration. Marine Biology 158,871-882.
Terrados J, Duarte CM, Kenworthy WJ (1997) Experimental evidence for apical dominance in the seagrass Cymodocea nodosa. Marine Ecology Progress Series 148,263-268.
Van Dijk JK, Waycott M, Van Tussenbroek BI, Ouborg J (2007) Polymorphic microsatellite markers for the Caribbean seagrass Thalassia testudinum Banks ex Konig. Molecular Ecology Notes 7,89-91.
Van Straalen NM, Roelofs D (2006) An Introduction to Ecological Genomics. Oxford University Press.
Vekemans X, Hardy OJ (2004) New insights from fine-scale spatial genetic structure analyses in plant populations. Molecular Ecology 13,921-935.
Vidondo B, Prairie YT, Blanco JM, Duarte CM (1997) Some aspects of the analysis of size spectra in aquatic ecology. Limnology and Oceanography 42,184-192.
Waycott M, Barnes PAG (2001) AFLP diversity within and between populations of the Caribbean seagrass Thalassia testudinum (Hydrocharitaceae). Marine Biology 139,1021-1028.
Waycott M, Duarte CM, Carruthers TJB, Orth RJ, Dennison WC, Olyarnik S, Calladine A, Fourqurean JW, Heck KL, Hughes AR, Kendrick GA, Kenworthy WJ, Short FT, Williams SL (2009) Accelerating loss of seagrasses across the globe threatens coastal ecosystems. Proceedings of the National Academy of Sciences of the United States of America 106, 12377-12381.
Waycott M, Procaccini G, Les DH, Reusch TBH (2006) Seagrass Evolution, Ecology And Conservation:A Genetic Perspective. In: Larkum, A.D.W., Orth R.J., Duarte, CM. (Eds.), Seagrass:biology, ecology and conservation, pp. PP.25-50. Springer, The Netherland.
Weir BS, Cockerham CC (1984) Estimating F-statistics for the analysis of population struture. Evolution 38,1358-1370.
White C, Selkoe KA, Watson J, Siegel DA, Zacherl DC, Toon en RJ (2010) Ocean currents help explain population genetic structure. Pwceedings of the Royal Society B:Biological Sciences 277,1685-1694.
Worm B, Barbier EB, Beaumont N, Duffy JE, Folke C, Halpern BS, Jackson JBC, Lotze HK, Micheli F, Palumbi SR, Sala E, Selkoe KA, Stachowicz JJ, Watson R (2006) Impacts of biodiversity loss on ocean ecosystem services. Science 314,787-790.
Wright S (1943) Isolation by distance. Genetics 28,114-138.
Wright S (1969) Evolution And The Genetics of Populations. In:The Theroy of Gene Frequencies, vol.2. University of Chicago Press.
Wyllie-Echeverria S, Talbot SL, Rearick JR (2010) Genetic structure and diversity of Zostera marina (Eelgrass) in the San Juan Archipelago, Washington, USA. Estuaries and Coasts 33,811-827.
Xu NN, Yu S, Zhang JG, Tsang PKE, Chen XY(2010) Microsatellite primers for Halophila ovalis and cross-amplification in H. Minor (Hydrocharitaceae). American Journal of Botany 97, E56-E57.
Yu S, Cui MY, Liu B, Wang XY Chen XY (2009) Development and characterization of microsatellite loci in Ruppia maritima L. (Ruppiaceae). Conseiration Genetics Resources 1.241.243.
Zane L,Bargelloni L,Patarnello T(2002)Strategies for microsatellite isolation:a review. Molecular Ecology 11,1-16.
范航清、彭胜、石雅君、郑杏雯(2007)广西北部湾沿海海草资源与研究状况.广西科学14289-295.
范航清、邱广龙、石雅君、李森(2011)中国亚热带海草生理生态学研究.科学出版社,北京,中国.
黄小平、黄良民(2007)中国南海海草研究.广东经济出版社,广州.
黄小平、黄良民、李颖虹、许战洲、方静威、黄道建、韩秋影、黄晖、谭烨辉、刘胜(2006)华南沿海主要海草床及其生境威胁.科学通报51,114-119.
黄小平、江志坚、张景平、施震、汪飞、叶丰、李磊(2010)广东沿海新发现的海草床.热带海洋学报29,132-135.
李强、万建民(2005)SSRHunter,一个本地化的SSR位点搜索软件的开发.遗传27,808-810.
苗苑、徐娜娜、于硕、耿艳、陈小勇(2009)海草克隆性及其种群遗传效应.生态学报29,3846-3853.
伍伯瑜(1991)黑潮和中国近海环流.台湾海峡10,25-32.
徐娜娜(2011)喜盐草的克隆性以及种群遗传效应与生态影响.华东师范大学,上海.
杨士瑛、鲍献文、陈长胜、陈菲(2003)夏季粤西沿岸流特征及其产生机制.海洋学报25,1-8.
俞慕耕、刘金芳(1993)南海海流系统与环流形势.海洋预报10,13-17.
于硕(2010)中国沿海川蔓藻(Ruppia)的分布及其影响因素.华东师范大学,上海.
张新宇、高燕宁(2004)PCR引物设计及软件使用技巧.生物信息学2,15-18.
张玉芬、张大勇(2004)克隆植物生态学与进化,In.植物生活史进化与繁殖生态学.科技出版社,北京.
Abe M, Yokota K, Kurashima A, Maegawa M (2009b) Temperature characteristics in seed germination and growth of Zostera japonica Ascherson & Graebner from Ago Bay, Mie Prefecture, central Japan. Fisheries Science 75,921-927.
Aioi K, Nakaoka M (2003) The Seagrass of Japan. In:Green EP and Short FT (eds) World Atlas of Seagrasses. University of California Press, Berkeley.
Alberto F, Correia L, Arnaud-Haond S, Billot C, Duarte CM, Serrao E (2003a) New microsatellite markers for the endemic Mediterranean seagrass Posidonia oceanica. Molecular Ecology Notes 3,253-255.
Alberto F, Correia L, Billot C, Duarte CM, Serrao E (2003b) Isolation and characterization of microsatellite markers for the seagrass Cymodocea nodosa. Molecular Ecology Notes 3, 397-399.
Alberto F, Gouveia L, Arnaud-Haond S, Perez-Llorens JL, Duarte CM, Serrao EA (2005) Within-population spatial genetic structure, neighbourhood size and clonal subrange in the seagrass Cymodocea nodosa. Molecular Ecology 14,2669-2681.
Alberto F, Massa S, Manent P, Diaz-Almela E, Arnaud-Haond S, Duarte CM, Serrao EA (2008) Genetic differentiation and secondary contact zone in the seagrass Cymodocea nodosa across the Mediterranean-Atlantic transition regions. Journal of Biogeography 35, 1279-1294.
Allendorf F W, Luikart G (2007) Conservation and the Genetics of Populations. Blackwell.
Anderson JB, Kohn LM (1995) Clonality in soilborne, plant-pathogenic fungi. Annual Review of Phytopathology 33,369-391.
Angevine MW, Handel SN (1986) Invasion of forest floor space, clonal architecture, and population growth in the perennial herb Clintonia Borealis. Journal of Ecology 74, 547-560.
Araki S, Kunii H (2006) Allozymic implications of the propagation of eelgrass Zostera japonica within a river system. Limnology 7,15-21.
Arnaud-Haond S, Alberto F, Teixeira S, Procaccini G, Serrao EA, Duarte CM (2005) Assessing genetic diversity in clonal organisms:low diversity or low resolution? Combining power and cost efficiency in selecting markers. Journal of Heredity 96,434-440.
Arnaud-Haond S, Belkhir K (2007) GENCLONE:a computer program to analyse genotypic data, test for clonality and describe spatial clonal organization. Molecular Ecology Notes 7, 15-17.
Arnaud-Haond S, Duarte CM, Alberto F, Serrao EA (2007a) Standardizing methods to address clonality in population studies. Molecular Ecology 16,5115-5139.
Arnaud-Haond S, Duarte CM, Diaz-Aimela E, Marba N, Sintes T, Sen-ao EA(2012) Implications of extreme life span in clonal organisms:millenary clones in meadows of the threatened seagrass Posidonia oceanica. PLoS ONE 7, e30454.
Arnaud-Haond S, Migliaccio M, Diaz-Almela E, Teixeira S, van de Vliet MS, Alberto F, Procaccini G, Duarte CM, Serrao EA (2007b) Vicariance patterns in the Mediterranean Sea:east-west cleavage and low dispersal in the endemic seagrass Posidonia oceanica. Journal of Biogeography 34,963-976.
Avise JC (2004) Molecular Markers, Natural History and Evolution. Sinauer Associates, Sunderland, Mass.
Ballesteros E, Cebrian E, Garcia-Rubies A, Alcoverro T, Romero J, Font X (2005) Pseudovivipary, a new fonn of asexual reproduction in the seagrass Posidonia oceanica. Botanica Marina 48,175-177.
Beaumont MA, Nichols RA(1996) Evaluating loci for use in the genetic analysis of population structure. Proceedings of the Royal Society of London Series B:Biological Sciences 263, 1619-1626.
Beebee T, Rowe G (2004) An Introduction to Molecular Ecology. Oxford University Press.
Beerli R (2011) Migrate-N, Version 3.2.17. http://popgen.scs.fsu.edu/mirate-n.html.
Bell AD, Tomlinson PB (1980) Adaptive architecture in rhizomatous plants. Botanical Journal of the Linnean Society 80,125-160.
Benjamin KJ, Walker DI, McComb AJ, Kuo J (1999) Structural response of marine and estuarine plants of Halophila ovalis (R-Br.) Hook. f. to long-term hyposalinity. Aquatic Botany 64, 1-17.
Billingham MR, Simoes T, Reusch TBH, Serrao EA(2007) Genetic sub-structure and intermediate optimal outcrossing distance in the marine angiosperm Zostera marina. Marine Biology 152,793-801.
Bohonak AJ (2002) IBD (Isolation by Distance):a program for analyses of isolation by distance. Journal of Heredity 93,153-154.
Brun FG, Cummaudo F, Olive I,Vergara JJ, Perez-Llorens JL (2007) Clonal extent, apical dominance and networking features in the phalanx angiosperm Zostera noltii Hornem. Marine Biology 151,1917-1927.
Cain ML, Dudle DA, Evans JP (1996) Spatial models of foraging in clonal plant species. American Journal of Botany 83,76-85.
Campbell SJ, McKenzie LJ, Kerville SP (2006) Photosynthetic responses of seven tropical seagrasses to elevated seawater temperature. Journal of Experimental Marine Biology and Ecology 330,455-468.
Capiomont A, Sandmeier M, Caye Q Meinesz A (1996) Enzyme polymorphism in Posidonia oceanica, a seagrass endemic to the Mediterranean. Aquatic Botany 54,265-277.
Casa AM, Mitchell SE, Hamblin MT, Sun H, Bowers JE, Paterson AH, Aquadro CF, Kresovich S (2005) Diversity and selection in sorghum:simultaneous analyses using simple sequence repeats. Theoretical and Applied Genetics 111,23-30.
Ceccherelli G, Campo D, Milazzo M (2007) Short-term response of the slow growing seagrass Posidonia oceanica to simulated anchor impact. Marine Environmental Research 63, 341-349.
Cornuet JM, Luikart G (1996) Description and power analysis of two tests for detecting recent population bottlenecks from allele frequency data. Genetics 144,2001-2014.
Coyer JA, Miller KA, Engle JM, Veldsink J, Cabello-Pasini A, Stam WT, Olsen JL (2008) Eelgrass meadows in the California channel islands and adjacent coast reveal a mosaic of two species, evidence for introgression and variable clonality. Annals of Botany 101, 73-87.
Coyer JA, Reusch TBH, Stam WT, Serrao EA, Pearson Q Procaccinis Q Olsen JL (2004) Characterization of microsatellite loci in the dwarf eelgrass Zostera noltii (Zosteraceae) and cross-reactivity with Z. japonica. Molecular Ecology Notes 4,497-499.
Cristescu R, Sherwin WB, Handasyde K, Cahill V, Cooper DW (2010) Detecting bottlenecks using BOTTLENECK 1.2.02 in wild populations:the importance of the microsatellite structure. Conservation Genetics 11,1043-1049.
Davis MB, Shaw RG (2001) Range shifts and adaptive responses to Quaternary climate change. Science 292,673-679.
Di Rienzo A, Peterson AC, Garza JC,'Valdes AM, Slatkin M, Freimer NB (1994) Mutational processes of simple-sequence repeat loci in human populations. Proceedings of the National Academy of Sciences of the United States of America 91,3166-3170.
Diaz-Almela E, Arnaud-Haond S, Vliet MS, Alvarez E, Marba N, Duarte CM, Serrao EA(2007) Feed-backs between genetic structure and perturbation-driven decline in seagrass (Posidonia oceanica) meadows. Conservation Genetics 8,1377-1391.
Diekmann OE, Coyer JA, Ferreira J, Olsen JL, Stam WT, Pearson GA, Serrao EA (2005) Popoulation genetics of Zostera noltii along the west Iberian coast:consequences of small population size, habitat discontinuity and near-shore currents. Marine Ecology Progress Series 290,89-96.
Dorken ME, Eckert CG (2001) Severely reduced sexual reproduction in northern populations of a clonal plant, Decodon verticillatus (Lythraceae). Journal of Ecology 89,339-350.
Douhovnikoff V, Dodd RS (2003) Intra-clonal variation and a similarity threshold for identification of clones:application to Salix exigua using AFLP molecular markers. Theoretical and Applied Genetics 106,1307-1315.
Duarte CM (1991) Allometric scaling of seagrass form and productivity. Marine Ecology Progress Series 77,289-300.
Duarte CM, Marba N, Krause-Jensen D, Sanchez-Camacho M (2007) Testing the predictive power of seagrass depth limit models. Estuaries andCoasts 30,652-656.
Epperson BK (1993) Recent advances in correlation studies of spatial patterns of genetic-variation. Evolutionary Biology 27,95-155.
Epperson BK (2003) Geographical Genetics:Monographs in Population Biology. Princeton University Press, Princeton, New Jersey.
Erftemeijer PLA, Stapel J (1999) Primary production of deep-water Halophila ovalis meadows. Aquatic Botany 65,71-82.
Evanno G, Regnaut S, Goudet J (2005) Detecting the number of clusters of individuals using the software STRUCTURE:a simulation study. Molecular Ecology 14,2611-2620.
Fenster CB,Vekemans X, Hardy OJ (2003) Quantifying gene flow from spatial genetic structure data in a metapopulation of Chamaecrista fasciculata (Leguminosae). Evolution 57, 995-1007.
Flint J, Bond J, Rees DC, Boyce AJ, Roberts-Thomson JM, Excoffier L, Clegg JB, Beaumont MA, Nichols RA, Harding RM (1999) Minisatellite mutational processes reduce Fst estimates. Human Genetics 105,567-576.
Gao H, Jiang K, Geng Y, Chen XY (2012) Development of microsatellite primers of the largest seagrass, Enhalus acoroides (Hydrocharitaceae). American Journal of Botany 99, E99-E101.
Gliddon C, Belhassen E, Gouyou PH (1987) Genetic neighborhoods in plants with diverse systems of mating and different patterns of growth. Heredity 59.29-32.
Goudet J (2002) FSTAT (Version 2.9.3.2):A computer program to calculate F-statistics. Updated from Goudet (1995). Available from http://www.unil.ch/izea/softwares/fstat.html.Journal of Heredity 86,485-486.
Halkett F, Simon JC, Balloux F (2005) Tackling the population genetics of clonal and partially clonal organisms. Trends in Ecology & Evolution 20,194-201.
Halpern BS, Walbridge S, Selkoe KA, Kappel CV, Micheli F, D'Agrosa C, Bruno JF, Casey KS, Ebert C, Fox HE, Fujita R, Heinemann D, Lenihan HS, Mad in EMP, Perry MT, Selig ER, Spalding M, Steneck R, Watson R (2008) A global map of human impact on marine ecosystems. Science 319,948-952.
Hammerli A, Reusch TBH (2003a) Genetic neighbourhood of clone structures in eelgrass meadows quantified by spatial autocorrelation of microsatellite markers. Heredity 91, 448-455.
Hammerli A, Reusch TBH (2003b) Inbreeding depression influences genet size distribution in a marine angiosperm. Molecular Ecology 12,619-629.
Harada Y, Iwasa Y(1996) Analyses of spatial patterns and population processes of clonal plants. Researches on Population Ecology (Kyoto) 38,153-164.
Hardy OJ, Vekemans X (1999) Isolation by distance in a continuous population:reconciliation between spatial autocorrelation analysis and population genetics models. Heredity 83, 145-154.
Hardy OJ, Vekemans X (2002) SPAGEDi:a versatile computer program to analyse spatial genetic structure at the individual or population levels. Molecular Ecology Notes 2,618-620.
Harrison PG, Bigley RE (1982) The recent introduction of the seagrass Zostera japonica Aschers. and Graebn. to the Pacific Coast of North America. Canadian Journal of Fisheries and Aquatic Sciences 39,1642-1648.
Hartog Cd, Kuo J (2006) Taxonomy and Biogeography of Seagrasses. In: Larkum, A.D.W., Orth, R.J., Duarte, CM. (Eds.), Seagrass:Biology, Ecology and Consei-vation. Springer, The Netherland.
Hedrick PW (2001) Conservation genetics:where are we now? Trends in Ecology & Evolution 16, 629-636.
Hedrick PW (2005) A standardized genetic differentiation measure. Evolution 59,1633-1638.
Hellberg ME (1995) Relationships between inferred levels of gene flow and geographic distance in a philopatric coral, Balanophyllia elegans. Evolution 48,1829-1854.
Hughes AR, Stachowicz JJ (2004) Genetic diversity enhances the resistance of a seagrass ecosystem to disturbance. Proceedings of the National Academy of Sciences of the United States of America 101,8998-9002.
Jiang K, Gao H, Xu NN, Tsang EPK, Chen XY(2011a) A set of microsatellite primers for Zostera japonica (Zosteraceae). American Journal of Botany 98, E236-E238.
Jiang K, Shi YS, Zhang J, Xu NN (2011b) Microsatellite primers for vulnerable seagrass Halophila beccarii (Hydrocharitaceae). American Journal of Botany 98, E155-E157.
Kendrick. GA, Duarte. CM, Marba. N (2005) Clonality in seagrasses, emergent properties and seagrass landscapes. Marine Ecology Progress Series 290,291-296.
Kornelis van Dijk J, van Tussenbroek BI (2010) Clonal diversity and structure related to habitat of the marine angiosperm Thalassia testudinum along the Atlantic coast of Mexico. Aquatic Botany 92,63-69.
Lalitha S (2000) Primer Primier 5. Biotech Software and Internet Report 1,270-273.
Lazrek F, Roussel V, Ronfort J, Cardinet Q Chardon F, Aouani M, Huguet T (2009) The use of neutral and non-neutral SSRs to analyse the genetic structure of a Tunisian collection of Medicago truncatula lines and to reveal associations with eco-environmental variables. Genetica 135,391-402.
Lee SY (1997) Annual cycle of biomass of a threatened population of the intertidal seagrass Zostera japonica in Hong Kong. Marine Biology 129,183-193.
Les DH, Cleland MA, Waycott M (1997) Phylogenetic studies in Alismatidae, Ⅱ:Evolution of marine angiosperms (seagrasses) and hydrophily. Systematic Botany 22,443-463.
Lirman D, Cropper WP (2003) The influence of salinity on seagrass growth, survivorship, and distribution within Biscayne Bay, Florida:Field, experimental, and modeling studies. Estuaries 26,131-141.
Lowe A, Harris S, Ashton P (2004) Ecological Genetics:Design, Analysis, and Application. Blackwell.
Lucas C, Thangaradjou T, Papenbrock J (2012) Development of a DNA barcoding system for seagrasses:successful but not simple. PLoSONE 7, e29987.
Manni F, Guerard E, Heyer E (2004) Geographic patterns of (genetic, morphologic, linguistic) variation:How barriers can be detected by using Monmonier's algorithm. Human Biology 76,173-190.
Marba N, Duarte CM (1998) Rhizome elongation and seagrass clonal growth. Marine Ecology Progress Series 174,269-280.
Marba N, Hemminga MA, Duarte CM (2006) Resource trans location within seagrass clones: allometric scaling to plant size and productivity. Oecologia 150,362-372.
Marba N, Hemminga MA, Mateo MA, Duarte CM, Mass YEM, Terrados J, Gacia E (2002) Carbon and nitrogen translocation between seagrass ramets. Marine Ecology Progress Series 226,287-300.
Meirmans PG (2006) Using the AMOVA framework to estimate a standardized genetic differentiation measure. Evolution 60,2399-2402.
Miller MP (1997) Tools for Population Genetic Analyses (TFPGA). A Windows program for the analysis of allozyme and molecular population genetic data version 1.3, Department of Biological Sciences, Northern Arizona University, Flagstaff, AZ.
Muniz-Salazar R, Talbot SL, Sage GK, Ward DH, Cabello-Pasini A (2005) Population genetic structure of annual and perennial populations of Zostera marina L. along the Pacific coast of Baja California and the Gulf of California. Molecular Ecology 14,711-722.
Nei M (1978) Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89,583-590.
Nielsen EE, Hansen MM, Meldrup D (2006) Evidence of microsatellite hitch-hiking selection in Atlantic cod (Gadus morhua L.):implications for inferring population structure in nonmodel organisms. Molecular Ecology 15,3219-3229.
Ohsawa T, Ide Y (2008) Global patterns of genetic variation in plant species along vertical and horizontal gradients on mountains. Global Ecology and Biogeography 17,152-163.
Peakall ROD, Smouse PE (2006) genalex 6:genetic analysis in Excel. Population genetic software for teaching and research. Molecular Ecology Notes 6,288-295.
Posey MH (1988) Community changes associated with the spread of an introduced seagrass, Zostera japonica. Ecology 69,974-983.
Prichard JK, Wen X, Falush D (2009) Documentation for structure software: version 2.3. http://pritch.bsd.uchicago.edu/structure.html.
Procaccini G, Mazzella L (1998) Population genetic structure and gene flow in the seagrass Posidonia oceanica assessed using microsatellite analysis. Marine Ecology Progress Series 169,133-141.
Procaccini G, Olsen JL, Reusch TBH (2007) Contribution of genetics and genomics to seagrass biology and conservation. Journal of Experimental Marine Biology and Ecology 350, 234-259.
Procaccini G, Orsini L, Ruggiero MV, Scardi M (2001) Spatial patterns of genetic diversity in Posidonia oceanica, an endemic Mediterranean seagrass. Molecular Ecology 10, 1413-1421.
Procaccini G, Piazzi L (2001) Genetic polymorphism and transplantation success in the mediterranean seagrass Posidonia oceanica. Restoration Ecology 9,332-338.
Procaccini G, Ruggiero MV, Orsini L (2002) Genetic structure and distribution of microsatellite population genetic diversity in Posidonia oceanica in the Mediterranean basin. Bulletin of Marine Science 71,1291-1297.
Procaccini G, Waycott M(1998) Microsatellite loci identified in the seagrass Posidonia oceanica (L.) Delile. Journal of Heredity 89,562-568.
Rannala B, Mountain JL (1997) Detecting immigration by using multilocus genotypes. Proceedings of the National Academy of Sciences of the United States of America 94, 9197-9201.
Reusch TBH (2000a) Five microsatellite loci in eelgrass Zostera marina and a test of cross-species amplification in Z. noltii and Z.japonica. Molecular Ecology 9,371-373.
Reusch TBH (2000b) Pollination in the marine realm:microsatellites reveal high outcrossing rates and multiple paternity in eelgrass Zostera marina. Heredity 85,459-464.
Reusch TBH (2001) New markers-old questions:population genetics of seagrasses. Marine Ecology Progress Series 211,261-274.
Reusch TBH, Ehlers A, Hammerli A, Worm B (2005) Ecosystem recovery after climatic extremes enhanced by genotypic diversity. Proceedings of the National Academy of Sciences of the United States of America 102,2826-2831.
Reusch TBH, Stam WT, Olsen JL (1999) Size and estimated age of genets in eelgrass, Zostera marina, assessed with microsatellite markers. Marine Biology 133,519-525.
Reusch TBH, Stam WT, Olsen JL(2000) Amicrosatellite-based estimation of clonal diversity and population subdivision in Zostera marina, a marine flowering plant. Molecular Ecology 9, 127-140.
Richardson CA, Kennedy H, Duarte CM. Kennedy DP, Proud SV (1999) Age and growth of the fan mussel Pinna nobilis from south-east Spanish Mediterranean seagrass (Posidonia oceanicu) meadows. Marine Biology 133,205-212.
Rousset F (1997) Genetic differentiation and estimation of gene flow from F-statistics under isolation by distance. Genetics 145,1219-1228.
Ruggiero MV, Capone S, Pirozzi P, Reusch TBH, Procaccini G (2005a) Mating system and clonal architecture: a comparative study in two marine angiosperms. Evolutionary Ecology 19, 487-499.
Ruggiero MV, Reusch TBH, Procaccini G (2004) Polymorphic microsatellite loci for the marine angiosperm Cymodocea nodosa. Molecular Ecology Notes 4,512-514.
Ruggiero MV, Reusch TBH, Procaccini G (2005b) Local genetic structure in a clonal dioecious angiosperm. Molecular Ecology 14,957-967.
Ruggiero MV, Turk R, Procaccini G (2002) Genetic identity and homozygosity in North-Adriatic populations of Posidonia oceanica:an ancient, post-glacial clone? Conservation Genetics 3,71-74.
Schlotterer C (2004) The evolution of molecular markers-just a matter of fashion? Nature Reviews Genetics 5,63-69.
Serra IA, Innocenti AM, Di Maida G, Calvo S, Migliaccio M, Zambianchi E, Pizzigalli C, Arnaud-Haond S, Duarte CM, Serrao EA, Procaccini G (2010) Genetic structure in the Mediterranean seagrass Posidonia oceanica:disentangling past vicariance events from contemporary patterns of gene flow. Molecular Ecology 19,557-568.
Serra IA, Procaccini G, Intrieri MC, Migliaccio M, Mazzuca S, Innocenti AM (2007) Comparison of lSSR and SSR markers for analysis of genetic diversity in the seagrass Posidonia oceanica. Marine Ecology Progress Series 338,71-79.
Shi MM, Michalski SG, Chen XY, Durka W (2011) Isolation by elevation:genetic structure at neutral and putatively non-neutral loci in a dominant tree of subtropical forests, Castanopsis eyrei. PLoSONE 6, e21302.
Shin H, Choi HK (1998) Taxonomy and distribution of Zostera (Zosteraceae) in eastern Asia, with special reference to Korea. Aquatic Botany 60,49-66.
Short F, Carruthers T, Dennison W, Waycott M (2007) Global seagrass distribution and diversity:a bioregional model. Journal of Experimental Marine Biology and Ecology 350,3-20.
Short FT, Polidoro B, Livingstone SR, Carpenter K.E, Bandeira S, Bujang JS, Calumpong HP, Carruthers TJB, Coles RG, Dennison WC, Erftemeijer PLA, Fortes MD, Freeman AS, Jagtap TG, Kamal AHM, Kendrick GA, Judson Kenworthy W, La Nafie YA, Nasution IM, Orth RJ, Prathep A, Sanciangco JC, Tussenbroek Bv, Vergara SG, Waycott M, Zieman JC (2011) Extinction risk assessment of the world's seagrass species. Biological Conservation 144,1961-1971.
Sinclair EA, Anthony J, Coupland GT, Waycott M, Barrett MD, Barrett RL, Cambridge ML, Wallace MJ, Dixon KW, Krauss SL, Kendrick GA (2009) Characterisation of polymorphic microsatellite markers in the widespread Australian seagrass, Posidonia australis Hook. f. (Posidoniaceae), with cross-amplification in the sympatric P. sinuosa. Conservation Genetics Resources 1,273-276.
Smouse PE, Peakall ROD, Gonzales EVA (2008) A heterogeneity test for fine-scale genetic structure. Molecular Ecology 17,3389-3400.
Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011) MEGA5:Molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28,2731-2739.
Tanaka N, Demise T, Ishii M, Shoji Y, Nakaoka M (2011) Genetic structure and gene flow of eelgrass Zostera marina populations in Tokyo Bay, Japan:implications for their restoration. Marine Biology 158,871-882.
Terrados J, Duarte CM, Kenworthy WJ (1997) Experimental evidence for apical dominance in the seagrass Cymodocea nodosa. Marine Ecology Progress Series 148,263-268.
Van Dijk JK, Waycott M, Van Tussenbroek BI, Ouborg J (2007) Polymorphic microsatellite markers for the Caribbean seagrass Thalassia testudinum Banks ex Konig. Molecular Ecology Notes 7,89-91.
Van Straalen NM, Roelofs D (2006) An Introduction to Ecological Genomics. Oxford University Press.
Vekemans X, Hardy OJ (2004) New insights from fine-scale spatial genetic structure analyses in plant populations. Molecular Ecology 13,921-935.
Vidondo B, Prairie YT, Blanco JM, Duarte CM (1997) Some aspects of the analysis of size spectra in aquatic ecology. Limnology and Oceanography 42,184-192.
Waycott M, Barnes PAG (2001) AFLP diversity within and between populations of the Caribbean seagrass Thalassia testudinum (Hydrocharitaceae). Marine Biology 139,1021-1028.
Waycott M, Duarte CM, Carruthers TJB, Orth RJ, Dennison WC, Olyarnik S, Calladine A, Fourqurean JW, Heck KL, Hughes AR, Kendrick GA, Kenworthy WJ, Short FT, Williams SL (2009) Accelerating loss of seagrasses across the globe threatens coastal ecosystems. Proceedings of the National Academy of Sciences of the United States of America 106, 12377-12381.
Waycott M, Procaccini G, Les DH, Reusch TBH (2006) Seagrass Evolution, Ecology And Conservation:A Genetic Perspective. In: Larkum, A.D.W., Orth R.J., Duarte, CM. (Eds.), Seagrass:biology, ecology and conservation, pp. PP.25-50. Springer, The Netherland.
Weir BS, Cockerham CC (1984) Estimating F-statistics for the analysis of population struture. Evolution 38,1358-1370.
White C, Selkoe KA, Watson J, Siegel DA, Zacherl DC, Toon en RJ (2010) Ocean currents help explain population genetic structure. Pwceedings of the Royal Society B:Biological Sciences 277,1685-1694.
Worm B, Barbier EB, Beaumont N, Duffy JE, Folke C, Halpern BS, Jackson JBC, Lotze HK, Micheli F, Palumbi SR, Sala E, Selkoe KA, Stachowicz JJ, Watson R (2006) Impacts of biodiversity loss on ocean ecosystem services. Science 314,787-790.
Wright S (1943) Isolation by distance. Genetics 28,114-138.
Wright S (1969) Evolution And The Genetics of Populations. In:The Theroy of Gene Frequencies, vol.2. University of Chicago Press.
Wyllie-Echeverria S, Talbot SL, Rearick JR (2010) Genetic structure and diversity of Zostera marina (Eelgrass) in the San Juan Archipelago, Washington, USA. Estuaries and Coasts 33,811-827.
Xu NN, Yu S, Zhang JG, Tsang PKE, Chen XY(2010) Microsatellite primers for Halophila ovalis and cross-amplification in H. Minor (Hydrocharitaceae). American Journal of Botany 97, E56-E57.
Yu S, Cui MY, Liu B, Wang XY Chen XY (2009) Development and characterization of microsatellite loci in Ruppia maritima L. (Ruppiaceae). Conseiration Genetics Resources 1.241.243.
Zane L,Bargelloni L,Patarnello T(2002)Strategies for microsatellite isolation:a review. Molecular Ecology 11,1-16.
范航清、彭胜、石雅君、郑杏雯(2007)广西北部湾沿海海草资源与研究状况.广西科学14289-295.
范航清、邱广龙、石雅君、李森(2011)中国亚热带海草生理生态学研究.科学出版社,北京,中国.
黄小平、黄良民(2007)中国南海海草研究.广东经济出版社,广州.
黄小平、黄良民、李颖虹、许战洲、方静威、黄道建、韩秋影、黄晖、谭烨辉、刘胜(2006)华南沿海主要海草床及其生境威胁.科学通报51,114-119.
黄小平、江志坚、张景平、施震、汪飞、叶丰、李磊(2010)广东沿海新发现的海草床.热带海洋学报29,132-135.
李强、万建民(2005)SSRHunter,一个本地化的SSR位点搜索软件的开发.遗传27,808-810.
苗苑、徐娜娜、于硕、耿艳、陈小勇(2009)海草克隆性及其种群遗传效应.生态学报29,3846-3853.
伍伯瑜(1991)黑潮和中国近海环流.台湾海峡10,25-32.
徐娜娜(2011)喜盐草的克隆性以及种群遗传效应与生态影响.华东师范大学,上海.
杨士瑛、鲍献文、陈长胜、陈菲(2003)夏季粤西沿岸流特征及其产生机制.海洋学报25,1-8.
俞慕耕、刘金芳(1993)南海海流系统与环流形势.海洋预报10,13-17.
于硕(2010)中国沿海川蔓藻(Ruppia)的分布及其影响因素.华东师范大学,上海.
张新宇、高燕宁(2004)PCR引物设计及软件使用技巧.生物信息学2,15-18.
张玉芬、张大勇(2004)克隆植物生态学与进化,In.植物生活史进化与繁殖生态学.科技出版社,北京.