<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 Spreadplants were planted in <str<strong>on</strong>g>the</str<strong>on</strong>g> field in <str<strong>on</strong>g>the</str<strong>on</strong>g> spring <str<strong>on</strong>g>of</str<strong>on</strong>g> 1981 into anarea <str<strong>on</strong>g>of</str<strong>on</strong>g> 1,000-1,300 m 2 . However, most <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Spartina saltmarsh <str<strong>on</strong>g>of</str<strong>on</strong>g> coastal China was developed from 0.5 kg <str<strong>on</strong>g>of</str<strong>on</strong>g> seedsfrom North Carolina.( Chung et al. 1985) Three morphs <str<strong>on</strong>g>of</str<strong>on</strong>g> S.alterniflora were identified by Xu & Zhou (1985) and Qin etal. (1985) based <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g>ir morphological data (culm height,leaf size, ear length and biomass), respectively, collectedfrom <str<strong>on</strong>g>the</str<strong>on</strong>g> garden <str<strong>on</strong>g>of</str<strong>on</strong>g> Nanjing University (Table 1) and fieldplots in Luoyuan Bay <str<strong>on</strong>g>of</str<strong>on</strong>g> East Sea (Table 2). Both <str<strong>on</strong>g>the</str<strong>on</strong>g>experiments showed that Georgia Spartina had <str<strong>on</strong>g>the</str<strong>on</strong>g> tallestand str<strong>on</strong>gest culms, and <str<strong>on</strong>g>the</str<strong>on</strong>g> highest biomass. Meanwhile,Chen and Chung (1991) reported that <str<strong>on</strong>g>the</str<strong>on</strong>g>re were threegenetic types in <str<strong>on</strong>g>the</str<strong>on</strong>g>ir studies <str<strong>on</strong>g>of</str<strong>on</strong>g> isozymes <str<strong>on</strong>g>of</str<strong>on</strong>g> EST, MDH andPOD <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> seedlings (Table 3). Since <str<strong>on</strong>g>the</str<strong>on</strong>g> people liked toplant higher and str<strong>on</strong>ger individuals <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> species, most <str<strong>on</strong>g>of</str<strong>on</strong>g><str<strong>on</strong>g>the</str<strong>on</strong>g> S. alterniflora plantati<strong>on</strong>s in China are now tall form(Georgia populati<strong>on</strong>) S. alterniflora. The tall form Spartinahad significant variati<strong>on</strong>s in survivorship, culm height andbiomass al<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g> elevati<strong>on</strong>al gradients <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Chinese coast(Chen & Chung 1991).IMPORTANT HUMAN FORCES DRIVING SPARTINAINVASIONSIn 1962, <str<strong>on</strong>g>the</str<strong>on</strong>g> Chinese Government decided to introduceSpartina from its native habitats for agricultural goalsaccording to <str<strong>on</strong>g>the</str<strong>on</strong>g> scientists’ recommendati<strong>on</strong>s. In 1963, S.anglica was introduced from Europe by <str<strong>on</strong>g>the</str<strong>on</strong>g> Chinesegovernment. In 1969, <str<strong>on</strong>g>the</str<strong>on</strong>g> first workshop was held by <str<strong>on</strong>g>the</str<strong>on</strong>g>government <str<strong>on</strong>g>of</str<strong>on</strong>g> Zhejiang Province to accelerate development<str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica marshes. From 1973 to 1978, severalworkshops were held by <str<strong>on</strong>g>the</str<strong>on</strong>g> governments <str<strong>on</strong>g>of</str<strong>on</strong>g> Jiangsu andShand<strong>on</strong>g provinces for <str<strong>on</strong>g>the</str<strong>on</strong>g> same purposes. Here we givesome <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> events that are associated with Spartina inchr<strong>on</strong>ological order:• 1978: Spartina research group <str<strong>on</strong>g>of</str<strong>on</strong>g> Nanjing University received<str<strong>on</strong>g>the</str<strong>on</strong>g> Nati<strong>on</strong>al Award for Natural Science <str<strong>on</strong>g>of</str<strong>on</strong>g>China.• 1978: The Institute <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina and Tideland Developmentat Nanjing University was founded by China’sMinistry <str<strong>on</strong>g>of</str<strong>on</strong>g> Educati<strong>on</strong>, toge<str<strong>on</strong>g>the</str<strong>on</strong>g>r with China’s Committee<str<strong>on</strong>g>of</str<strong>on</strong>g> Science and Technology, China’s Ministry <str<strong>on</strong>g>of</str<strong>on</strong>g> Agricultureand China’s Ocean Bureau. The institute c<strong>on</strong>sisted<str<strong>on</strong>g>of</str<strong>on</strong>g> 17 members.• 1978-1979: Two workshops were held by China’sCommittee <str<strong>on</strong>g>of</str<strong>on</strong>g> Science and Technology to accelerate introducti<strong>on</strong><str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglica.• 1979: Three o<str<strong>on</strong>g>the</str<strong>on</strong>g>r Spartina species were introduced byPr<str<strong>on</strong>g>of</str<strong>on</strong>g>essor Chung-Hsin Chung, funded by <str<strong>on</strong>g>the</str<strong>on</strong>g> Chinesegovernment.• 1985: The first Chinese Spartina research m<strong>on</strong>ograph“Research Advances in Spartina — Achievement <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>Past 22 Years”, was published as a special issue <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>Journal <str<strong>on</strong>g>of</str<strong>on</strong>g> Nanjing University; <str<strong>on</strong>g>the</str<strong>on</strong>g> m<strong>on</strong>ograph c<strong>on</strong>tainedthree reviews, 26 reports and 24 short communicati<strong>on</strong>s.• 1985-1990: Several h<strong>on</strong>ors were awarded to Pr<str<strong>on</strong>g>of</str<strong>on</strong>g>.Chung by different Chinese government departments forhis distinguished achievement in introducti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>Spartina.• 1992: A m<strong>on</strong>ograph, Applied Studies <strong>on</strong> Spartina (eds.Qin, P. and C.H. Chung), was published by OceanPress; it c<strong>on</strong>tained 30 papers.• 1993: Pr<str<strong>on</strong>g>of</str<strong>on</strong>g>. Chung received an award from <str<strong>on</strong>g>the</str<strong>on</strong>g> Society<str<strong>on</strong>g>of</str<strong>on</strong>g> Wetland Scientists.• 1995: Pr<str<strong>on</strong>g>of</str<strong>on</strong>g>. Chung received a Distinguished FellowAward from Ohio State University.• 1996: The Spartina Research Group <str<strong>on</strong>g>of</str<strong>on</strong>g> Nanjing Universityreceived a Distinguished C<strong>on</strong>tributi<strong>on</strong> Award from<str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> Society <str<strong>on</strong>g>of</str<strong>on</strong>g> Ecological Engineering.• 1998: S. patens was again introduced into China as asalt-tolerant ec<strong>on</strong>omic species.CURRENT STATUS OF SPARTINA INVASIONS IN CHINAIn 1985, S. anglica plantati<strong>on</strong>, with human aid, reachedto an area <str<strong>on</strong>g>of</str<strong>on</strong>g> 36,000 ha, and was distributed in 83 coastalcounties in China. The species’ range had increased by 330times from 1966 to 1985. Most <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> plants came from 21English individuals. After that time, however, <str<strong>on</strong>g>the</str<strong>on</strong>g> specieshas declined without human plantings. Only 50 ha <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 still existed in 2000 (Table 4). The species hasbeen also experiencing dieback because it cannot produceTable 4. Area changes and <str<strong>on</strong>g>the</str<strong>on</strong>g>ir causes <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina in China. (Partialdata from Chung et al. 1985.)Year Spartina anglicaArea Causes(ha)1966 110 Planting and naturalreproducti<strong>on</strong>1973 2,000 Planting and naturalreproducti<strong>on</strong>1978 10,295 Planting and naturalreproducti<strong>on</strong>1980 31,590 Planting and naturalreproducti<strong>on</strong>1985 36,000 Planting and naturalreproducti<strong>on</strong>Area(ha)Spartina alternifloraCauses0.13 Planting260 Planting and naturalreproducti<strong>on</strong>2000
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> Spartinaany seeds and it has become shorter at all elevati<strong>on</strong>s.On <str<strong>on</strong>g>the</str<strong>on</strong>g> o<str<strong>on</strong>g>the</str<strong>on</strong>g>r hand, <str<strong>on</strong>g>the</str<strong>on</strong>g> range <str<strong>on</strong>g>of</str<strong>on</strong>g> S. alterniflora increasedby 10,000 times from 1980 to 1988. Alhough planting <str<strong>on</strong>g>the</str<strong>on</strong>g>species <strong>on</strong> a large scale was stopped in 1995, it stillincreased by 86 times from 1988 to 2000. The species is stillinvading coastal areas in China by natural dispersal and hasbecome a threat to <str<strong>on</strong>g>the</str<strong>on</strong>g> native ecosystems. From 1995 to2000, ca. 200,000 ha <str<strong>on</strong>g>of</str<strong>on</strong>g> S. alterniflora marshes and tidelandswere reclaimed, which did not stop <str<strong>on</strong>g>the</str<strong>on</strong>g> invasi<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.Many native species, including plants, some endangeredbirds (Ma et al. 2004), and mollusks <str<strong>on</strong>g>of</str<strong>on</strong>g> ec<strong>on</strong>omic value incoastal areas are threatened by S. alterniflora invasi<strong>on</strong>s. Itexcluded almost all <str<strong>on</strong>g>the</str<strong>on</strong>g> o<str<strong>on</strong>g>the</str<strong>on</strong>g>r native plants that wereoriginally dominant in wetlands, including Phragmitesaustralis, Typha spp., Scirpus spp., Suadae spp., and eveninvaded fishp<strong>on</strong>ds and young mangrove swamps (Qian andMa 1995). For example, in intertidal z<strong>on</strong>es <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> YangziRiver estuary, S. alterniflora invaded Scirpus mariquetercommunities. Through competiti<strong>on</strong> with <str<strong>on</strong>g>the</str<strong>on</strong>g> native species,S. alterniflora has greatly decreased native speciesabundance, and even excluded <str<strong>on</strong>g>the</str<strong>on</strong>g>m (Li et al. in thisvolume). Li et al. (this volume) also compared <str<strong>on</strong>g>the</str<strong>on</strong>g> structure<str<strong>on</strong>g>of</str<strong>on</strong>g> nematode communities am<strong>on</strong>g Spartina marsh and S.mariqueter and Phragmites marshes <str<strong>on</strong>g>of</str<strong>on</strong>g> D<strong>on</strong>gtan wetlands <strong>on</strong>Ch<strong>on</strong>gming Island, and found significant differences introphic structure <str<strong>on</strong>g>of</str<strong>on</strong>g> nematode communities between <str<strong>on</strong>g>the</str<strong>on</strong>g>marshes.Spartina alterniflora is <strong>on</strong>e <str<strong>on</strong>g>of</str<strong>on</strong>g> 16 notorious invasivepest plants in China as <str<strong>on</strong>g>the</str<strong>on</strong>g> species directly causes milli<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g>dollars <str<strong>on</strong>g>of</str<strong>on</strong>g> ec<strong>on</strong>omic loss per year (An et al, 2007). Spartinaalterniflora is still rapidly spreading, although <str<strong>on</strong>g>the</str<strong>on</strong>g> Chinesegovernment and scientists are doing <str<strong>on</strong>g>the</str<strong>on</strong>g>ir best to c<strong>on</strong>trol oreradiate <str<strong>on</strong>g>the</str<strong>on</strong>g> species by physical, chemical, biological andintegrated methods (Lin 1997; Liu and Huang 2000).RESEARCH BEFORE 1995Since S. anglica arrived at Nanjing University <str<strong>on</strong>g>of</str<strong>on</strong>g> China,a series <str<strong>on</strong>g>of</str<strong>on</strong>g> studies (all <str<strong>on</strong>g>the</str<strong>on</strong>g> papers published before 1986 in<str<strong>on</strong>g>the</str<strong>on</strong>g> cited literature) had been carried out by <str<strong>on</strong>g>the</str<strong>on</strong>g> SpartinaResearch Group, which focused <strong>on</strong> S. anglica by 1985. Forexample:• 1963 to 1966: Asexual propagati<strong>on</strong> from rhizome andramets; seed germinati<strong>on</strong>, tolerance <str<strong>on</strong>g>of</str<strong>on</strong>g> individuals tocold, hot, drought, water logging, silt sediment and saltstress; growth under different elevati<strong>on</strong>; sexual andasexual reproducti<strong>on</strong> from established individuals;transplanting and management techniques.• 1967 to 1973: Field transplanting; impacts <str<strong>on</strong>g>of</str<strong>on</strong>g> S. anglicagrowth <strong>on</strong> soil properties and planting techniques ininland riparian wetlands.• 1974 to 1978: Green manure for cropland, forage forsheep, goat and pig; height growth; fertilized flowersTable 5. Survivorship and growth <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina alterniflora at differentelevati<strong>on</strong> levels. (Revised from Xu and Zhuo 1985.)Elevati<strong>on</strong>(m)Inundati<strong>on</strong>frequency(days/m<strong>on</strong>th)Inundati<strong>on</strong>time(hours/day)Survivorship(%)Increase rate <str<strong>on</strong>g>of</str<strong>on</strong>g>individuals perclump (times)and seed producti<strong>on</strong>; biomass producti<strong>on</strong>, net biomassproducti<strong>on</strong>; impacts <str<strong>on</strong>g>of</str<strong>on</strong>g> harvest <strong>on</strong> growth and reproducti<strong>on</strong>and impacts <str<strong>on</strong>g>of</str<strong>on</strong>g> grazing by cows, sheep and goats.• 1979 to 1985: Silt sediment rate <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina plantati<strong>on</strong>;uses in protecti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> dykes; preventi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> tideland erosi<strong>on</strong>;purificati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> polluted water; impact <strong>on</strong> beneathfauna; Gram negative bacteria compositi<strong>on</strong>; anatomy <str<strong>on</strong>g>of</str<strong>on</strong>g>roots and stems; germinati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> seeds produced inChina and impacts <str<strong>on</strong>g>of</str<strong>on</strong>g> puncture and air pressure <strong>on</strong> seedgerminati<strong>on</strong>; physiology; tissue culture; introducti<strong>on</strong> toriparian wetlands in Yellow River <str<strong>on</strong>g>of</str<strong>on</strong>g> NorthwesternChina; biochemical compositi<strong>on</strong> and nutrient; impacts<str<strong>on</strong>g>of</str<strong>on</strong>g> fertilizer <strong>on</strong> growth and biomass producti<strong>on</strong>; foragefor fish; structures <str<strong>on</strong>g>of</str<strong>on</strong>g> salt glands; ultra-structures <str<strong>on</strong>g>of</str<strong>on</strong>g> leafcells; DNA c<strong>on</strong>tents; ecotype identificati<strong>on</strong> and methodsto increase <str<strong>on</strong>g>the</str<strong>on</strong>g> seed producti<strong>on</strong>.After 1985, most studies (in <str<strong>on</strong>g>the</str<strong>on</strong>g> cited literature between1986 and 1994) focused <strong>on</strong> biology and ecology <str<strong>on</strong>g>of</str<strong>on</strong>g> S.alterniflora, including survivorship and growth at differentelevati<strong>on</strong>s (Table 5); growth rate and growth form/type fromthree different origins; above- and below-ground biomassand biomass allocati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> three growth-type populati<strong>on</strong>s;seed germinati<strong>on</strong> at different saline stresses; effect <str<strong>on</strong>g>of</str<strong>on</strong>g> N-fixed 4088 strain <str<strong>on</strong>g>of</str<strong>on</strong>g> bacteria <strong>on</strong> seed germinati<strong>on</strong> andindividual growth; transportati<strong>on</strong> and distributi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>phosphorus (P) in organs under salt stress; physiology andbiochemistry <str<strong>on</strong>g>of</str<strong>on</strong>g> leaf and root; effects <str<strong>on</strong>g>of</str<strong>on</strong>g> seed soaking <strong>on</strong>seed germinati<strong>on</strong>; enzyme activity, sugar c<strong>on</strong>tents, aminoacid and cold tolerance <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> seedling under different saltstress; translocati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> mineral elements; c<strong>on</strong>tents, allocati<strong>on</strong>and uses <str<strong>on</strong>g>of</str<strong>on</strong>g> Selenium (Se) <str<strong>on</strong>g>of</str<strong>on</strong>g> plants; flav<strong>on</strong>oids andimmunity activity; isozyme differentiati<strong>on</strong>; micromorphology<str<strong>on</strong>g>of</str<strong>on</strong>g> pollen grains; ultrastructure <str<strong>on</strong>g>of</str<strong>on</strong>g> mesophyllcells; anatomy <str<strong>on</strong>g>of</str<strong>on</strong>g> seeds; biomass dynamics, speciesstructures, energy storage and energy flow <str<strong>on</strong>g>of</str<strong>on</strong>g> ecosystem; <str<strong>on</strong>g>the</str<strong>on</strong>g>relati<strong>on</strong>ships between Spartina and clamworms; soil enzymeactivity in Spartina salt marsh; extracti<strong>on</strong> method,toxicology, nutrient c<strong>on</strong>tents, functi<strong>on</strong>s and uses <str<strong>on</strong>g>of</str<strong>on</strong>g>biomineral beverages/tea.Height(cm)0.0 30 5.0 - 8.0 16.7 0 12.50.5 30 3.5 - 7.0 66.7 1.5 27.61.0 30 1.5 - 6.0 66.7 4.8 37.31.5 30 0.5 - 4.5 100.0 10.8 90.22.0 26 - 30 0.5 - 3.5 100.0 13.7 101.42.5 18 - 28 0.5 - 2.0 100.0 11.8 98.23.0 14 - 22 0.5 - 2.0 100.0 8.0 72.3-68-
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FORWARD & ACKNOWLEDGEMENTSThe <stro
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CHAPTER ONESpartina Biology
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Chapter 1: Spartina Biology
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