<str<strong>on</strong>g>Proceedings</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Third</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> <str<strong>on</strong>g>C<strong>on</strong>ference</str<strong>on</strong>g> <strong>on</strong> <strong>Invasive</strong> SpartinaChapter 2: Spartina Distributi<strong>on</strong> and SpreadSPARTINA IN CHINA:INTRODUCTION,HISTORY,CURRENT STATUS, AND RECENT RESEARCHS. AN,H.QING,Y.XIAO,C.ZHOU,Z.WANG,Z.DENG,Y.ZHI AND L. CHENSchool <str<strong>on</strong>g>of</str<strong>on</strong>g> Life Science, Nanjing University, Nanjing 210093, China; anshq@nju.edu.cnSpartina spp., with <str<strong>on</strong>g>the</str<strong>on</strong>g>ir str<strong>on</strong>g ability for survival, growth, and expansi<strong>on</strong>, attracts ecologists’attenti<strong>on</strong> throughout <str<strong>on</strong>g>the</str<strong>on</strong>g> world. Four Spartina species: Spartina anglica, S. alterniflora, S. patensand S. cynosuroides were introduced into China in 1963, 1979, 1979 and 1998, respectively, for <str<strong>on</strong>g>the</str<strong>on</strong>g>purposes <str<strong>on</strong>g>of</str<strong>on</strong>g> agricultural and ecological engineering. So far <str<strong>on</strong>g>the</str<strong>on</strong>g> first three species still survive incoastal China with varying abundance. Spartina anglica is dying back, whereas S. alterniflora hasbeen rampantly invading. Spartina patens appears to be a potential invasive species in China. Weexamine <str<strong>on</strong>g>the</str<strong>on</strong>g> history <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina’s introducti<strong>on</strong>s into China, discuss <str<strong>on</strong>g>the</str<strong>on</strong>g> fate <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> introduced speciesand <str<strong>on</strong>g>the</str<strong>on</strong>g>ir impacts <strong>on</strong> native ecosystems, and review <str<strong>on</strong>g>the</str<strong>on</strong>g> studies c<strong>on</strong>ducted in China over <str<strong>on</strong>g>the</str<strong>on</strong>g> past 40years (from 1963-2003), especially those carried out in <str<strong>on</strong>g>the</str<strong>on</strong>g> Institute <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina and TidelandDevelopment at Nanjing University.Keywords: Ecological engineering, genetic structure, invasive impacts, management, SpartinaINTRODUCTIONFour Spartina species have been introduced into Chinaas bio-engineers for agricultural and ecological engineeringsince 1963. They were used to accelerate <str<strong>on</strong>g>the</str<strong>on</strong>g> development<str<strong>on</strong>g>of</str<strong>on</strong>g> coastal tidelands for croplands, protect <str<strong>on</strong>g>the</str<strong>on</strong>g> dykes fromtypho<strong>on</strong>s and c<strong>on</strong>trol erosi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> tidelands from tidal waves(Chung and Zhuo 1979; Chung 1982, 1985). The specieshave had different fates in coastal areas <str<strong>on</strong>g>of</str<strong>on</strong>g> China resultingfrom both human activities and natural stresses. Seeds <str<strong>on</strong>g>of</str<strong>on</strong>g> S.patens and S. cynosuroides were introduced into China,toge<str<strong>on</strong>g>the</str<strong>on</strong>g>r with S. alterniflora. But, S. patens failed togerminate in <str<strong>on</strong>g>the</str<strong>on</strong>g> laboratory and S. cynosuroides failed tosurvive in <str<strong>on</strong>g>the</str<strong>on</strong>g> field. In 1998, seeds <str<strong>on</strong>g>of</str<strong>on</strong>g> S. patens wereintroduced into China again and successfully germinatedafter i<strong>on</strong> radiati<strong>on</strong> treatment (Zhou 2003), <str<strong>on</strong>g>the</str<strong>on</strong>g>n tissue culturewas used to produce <str<strong>on</strong>g>of</str<strong>on</strong>g>fspring at a large scale. So far, about20 hectares (ha) <str<strong>on</strong>g>of</str<strong>on</strong>g> S. patens has been transplanted inTianjing, Jiangsu and Zhejiang Provinces (Zhou et al 2003).Spartina alterniflora has rapidly spread to o<str<strong>on</strong>g>the</str<strong>on</strong>g>r coastalareas in China, outcompeted native plants and become <strong>on</strong>e<str<strong>on</strong>g>of</str<strong>on</strong>g> most harmful invading plants in China. In c<strong>on</strong>trast, S.anglica is experiencing a dieback and may disappear from<str<strong>on</strong>g>the</str<strong>on</strong>g> coasts. Although S. anglica is a serious invasive speciesin o<str<strong>on</strong>g>the</str<strong>on</strong>g>r countries (Daehler and Str<strong>on</strong>g 1996; Kriworken andHedge 2000; Baumel et al. 2001; Hacker et al. 2001), it isnow not an invasive species in China. More recently, S.patens has shown <str<strong>on</strong>g>the</str<strong>on</strong>g> potential to be an invasive species(Zhou et al. 2003). The four species have had differentdifferent fates including <str<strong>on</strong>g>the</str<strong>on</strong>g> failure <str<strong>on</strong>g>of</str<strong>on</strong>g> S. cynosuroidesintroducti<strong>on</strong> and different invasive processes in coastalChina. Here we give an brief account <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> experiencesfrom Chinese scientists and studies that have been c<strong>on</strong>ductedin China since <str<strong>on</strong>g>the</str<strong>on</strong>g> introducti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>se species to thiscountry. It is hoped that <str<strong>on</strong>g>the</str<strong>on</strong>g> informati<strong>on</strong> presented here canbe <str<strong>on</strong>g>of</str<strong>on</strong>g> some use to <str<strong>on</strong>g>the</str<strong>on</strong>g> scientists who are working <strong>on</strong> use,management, c<strong>on</strong>trol and eradicati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> species that areinvading both Atlantic and Pacific coastal areas worldwide.HISTORY OF SPARTINA INTRODUCTION IN CHINASpartina anglicaSpartina anglica was <str<strong>on</strong>g>the</str<strong>on</strong>g> first Spartina speciesintroduced into China from ano<str<strong>on</strong>g>the</str<strong>on</strong>g>r country. In 1963, 35plants from Essex in England and 100 plants from Hojer inDenmark were introduced to <str<strong>on</strong>g>the</str<strong>on</strong>g> ecological laboratory <str<strong>on</strong>g>of</str<strong>on</strong>g>Nanjing University by Pr<str<strong>on</strong>g>of</str<strong>on</strong>g>. Chung-Hsin Chung (Ch<strong>on</strong>g-XinZh<strong>on</strong>g) with <str<strong>on</strong>g>the</str<strong>on</strong>g> help <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Chinese Committee <str<strong>on</strong>g>of</str<strong>on</strong>g> Scienceand Technology. The Spartina survivors — 21 from Englandand 50 from Denmark — were sent to SheyangExperimental Stati<strong>on</strong> (35 o N) <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> Yellow Sea coast after asimple survivial check in <str<strong>on</strong>g>the</str<strong>on</strong>g> laboratory. The survivors <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>English Spartina successfully produced 435,000 new rametsfrom July 1963 to April 1964, which were planted in <str<strong>on</strong>g>the</str<strong>on</strong>g>field in 1964; by 1966 <str<strong>on</strong>g>the</str<strong>on</strong>g> area <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> plantati<strong>on</strong> hadincreased to 32 ha. However, <str<strong>on</strong>g>the</str<strong>on</strong>g> Danish populati<strong>on</strong> so<strong>on</strong>died <str<strong>on</strong>g>of</str<strong>on</strong>g>f. Almost all <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> S. anglica in China aredescendants <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> original 21 English plants, excluding <str<strong>on</strong>g>the</str<strong>on</strong>g>plantati<strong>on</strong>s in Xiaoshan and Wenlin <str<strong>on</strong>g>of</str<strong>on</strong>g> Zhejiang Province. In1964, 507 seeds and 18 plants from Poole Harbor in Englandwere introduced to China, <str<strong>on</strong>g>of</str<strong>on</strong>g> which 440 seeds with potentialgerminati<strong>on</strong> capacity were treated and sowed in <str<strong>on</strong>g>the</str<strong>on</strong>g>laboratory; 157 <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>m germinated, a germinati<strong>on</strong> rate <str<strong>on</strong>g>of</str<strong>on</strong>g>30.97%, and 44 seedlings survived. The 18 plants died <str<strong>on</strong>g>of</str<strong>on</strong>g>f in<str<strong>on</strong>g>the</str<strong>on</strong>g> lab. One year later, 30,601 individuals were obtainedfrom <str<strong>on</strong>g>the</str<strong>on</strong>g> 44 surviving seedlings (Fig. 1), and most <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>mwere planted in Xiaoshan and Wenlin Experimental Stati<strong>on</strong>s-65-
Chapter 2: Spartina Distributi<strong>on</strong> and Spread<str<strong>on</strong>g>Proceedings</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Third</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> <str<strong>on</strong>g>C<strong>on</strong>ference</str<strong>on</strong>g> <strong>on</strong> <strong>Invasive</strong> Spartinaal<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g> coast <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> East Sea (29 o N). In <str<strong>on</strong>g>the</str<strong>on</strong>g> experiment,<strong>on</strong>e rhizome segment produced 9.10 milli<strong>on</strong> individualsduring 29 m<strong>on</strong>ths, and four ramets produced 7.60 milli<strong>on</strong>individuals during 33 m<strong>on</strong>ths. Most <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> 16.70 milli<strong>on</strong>individuals were planted in mid-1966 in two East Seastati<strong>on</strong>s, whose area was 73 ha. The total area <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglicaplantati<strong>on</strong> was 78 ha by August 1966 in <str<strong>on</strong>g>the</str<strong>on</strong>g> two stati<strong>on</strong>s.But, from 1973 to 1980, most <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> plantati<strong>on</strong>s in <str<strong>on</strong>g>the</str<strong>on</strong>g> twostati<strong>on</strong>s were reclaimed for croplands; thus almost all <str<strong>on</strong>g>the</str<strong>on</strong>g>individuals <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica died <str<strong>on</strong>g>of</str<strong>on</strong>g>f <str<strong>on</strong>g>the</str<strong>on</strong>g>re.Four Spartina species: Spartina anglica, S. alterniflora,S. patens and S. cynosuroides have been introduced intoChina in 1963, 1979, 1979 and 1998, respectively, for <str<strong>on</strong>g>the</str<strong>on</strong>g>purposes <str<strong>on</strong>g>of</str<strong>on</strong>g> agricultural and ecological engineering.By 1978,all <str<strong>on</strong>g>the</str<strong>on</strong>g> coastal provinces <str<strong>on</strong>g>of</str<strong>on</strong>g> China planted S. anglica. Therewere 6,330 ha, 3,750 ha and 200 ha <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica plantati<strong>on</strong>sin Jiangsu, Zhejiang and Shand<strong>on</strong>g Provinces, respectively.Meanwhile, 5 ha, 2/3 ha, 1/2 ha and 1/3 ha existed in Hebei,Lia<strong>on</strong>ing, Tianjing and Guangd<strong>on</strong>g Provinces, respectively.Less than 1/3 ha <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> species was established in Shanghai,Fujian and Guangxi Provinces. In <str<strong>on</strong>g>the</str<strong>on</strong>g> field, S. anglica had astr<strong>on</strong>g reproductive capacity by asexual propagati<strong>on</strong> throughramets and rhizomes (Fig. 2). By 1980, all 83 counties <str<strong>on</strong>g>of</str<strong>on</strong>g>coastal China had S. anglica with a total area <str<strong>on</strong>g>of</str<strong>on</strong>g> 31,590 ha;by 1985 area had increased to 36,000 ha, although <str<strong>on</strong>g>the</str<strong>on</strong>g> sexualreproducti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> species was very low.Spartina alternifloraSimilarly, seeds and individuals <str<strong>on</strong>g>of</str<strong>on</strong>g> S. alterniflora werealso introduced to China by Pr<str<strong>on</strong>g>of</str<strong>on</strong>g>essor Chung at <str<strong>on</strong>g>the</str<strong>on</strong>g> end <str<strong>on</strong>g>of</str<strong>on</strong>g>1979 from North Carolina, Georgia and Florida, USA.Seedlings were successfully obtained from <str<strong>on</strong>g>the</str<strong>on</strong>g> seeds andnew ramets were reproduced from <str<strong>on</strong>g>the</str<strong>on</strong>g> exotic individuals in<str<strong>on</strong>g>the</str<strong>on</strong>g> laboratory and garden <str<strong>on</strong>g>of</str<strong>on</strong>g> Nanjing University in 1980,and <str<strong>on</strong>g>the</str<strong>on</strong>g>n were sent to Luoyuanwan Stati<strong>on</strong> (26 0 30’). TheTable 1. Growth <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>on</strong>e-year Spartina alterniflora from three differentorigins. (Data from Xu & Zhuo 1985.)Culm height(cm)Leaf length(cm)Leaf width(cm)Ear length(cm)SourcesRange Max Range Max Range Max Range MaxBiomass(DW g/m 2)LeafcolorNorthCarolina 110-170 217 45-70 95.5 1.3-1.5 2.0 14-30 48 297.9 BlackgreenLightGeorgia 140-240 275 50-70 90.0 1.4-1.7 2.1 18-35 42 457.2greenFlorida 70-100 128 40-60 82.0 1.5-1.7 2.1 12-15 30 268.2 GreenTable 2. Height and biomass growth <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina alterniflora fromdifferent origins at <str<strong>on</strong>g>the</str<strong>on</strong>g> field plots (Xu & Zhuo 1985).Fig. 1 Asexual reproducti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> exotic Spartina anglica from seedlings.(Data from Chung et al 1985.)IntroductiveSourcesCulmheight(cm)Above-groundbiomass(g DW/m 2)Below-groundbiomass(g DW/m 2)Total biomass(g DW/m 2)North Carolina 170 (35) 1289 (136) 1235 (160) 2524 (274)Georgia 240 (50) 2745 (502) 1366 (25) 4111 (486)Florida 110 (25) 972 (139) 928 (100) 1900 (83)ANOVA & T-testP
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CHAPTER ONESpartina Biology
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