<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 4: Spartina C<strong>on</strong>trol and ManagementBIOLOGICAL CONTROL OF SPARTINAF. S. GREVSTAD 1 ,M.S.WECKER 1 , AND D. R. STRONG 21 Olympic Natural Resources Center, University <str<strong>on</strong>g>of</str<strong>on</strong>g> Washingt<strong>on</strong>, P.O. Box 1628, Forks, WA 98331;grevstad@u.washingt<strong>on</strong>.edu2 Department <str<strong>on</strong>g>of</str<strong>on</strong>g> Evoloti<strong>on</strong> and Ecology, University <str<strong>on</strong>g>of</str<strong>on</strong>g> California, Davis, CA 95616Biological c<strong>on</strong>trol using introduced natural enemies can be an effective approach to <str<strong>on</strong>g>the</str<strong>on</strong>g> l<strong>on</strong>g termc<strong>on</strong>trol <str<strong>on</strong>g>of</str<strong>on</strong>g> widespread weeds. A biological c<strong>on</strong>trol program against Spartina spp. is underway inWashingt<strong>on</strong> State, where more than 10,000 hectares (ha) <str<strong>on</strong>g>of</str<strong>on</strong>g> intertidal mudflat are affected bySpartina alterniflora and Spartina anglica. Releases <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> planthopper Prokelisia marginata havebeen made into Willapa Bay each year since 2000 and into Puget Sound since 2003. Prior tointroducing this insect, rigorous host specificity testing and a review by <str<strong>on</strong>g>the</str<strong>on</strong>g> Technical AdvisoryGroup <strong>on</strong> Biological C<strong>on</strong>trol <str<strong>on</strong>g>of</str<strong>on</strong>g> Weeds c<strong>on</strong>firmed that <str<strong>on</strong>g>the</str<strong>on</strong>g> risk to n<strong>on</strong>-target plants was minute.Populati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> bioc<strong>on</strong>trol agent were initially slow to establish and grow. However, earlyproblems with high winter mortality have been remedied through a combinati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> improved releasesite selecti<strong>on</strong> and <str<strong>on</strong>g>the</str<strong>on</strong>g> use <str<strong>on</strong>g>of</str<strong>on</strong>g> cold-hardy east coast biotypes. At least two populati<strong>on</strong>s in Willapa Bayare well established and expanding. At a localized scale, we have measured 50 percent reducti<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g>Spartina biomass and 90 percent reducti<strong>on</strong> in viable seed set due to P. marginata. The full extent <str<strong>on</strong>g>of</str<strong>on</strong>g><str<strong>on</strong>g>the</str<strong>on</strong>g> impact will <strong>on</strong>ly be known with time.While <str<strong>on</strong>g>the</str<strong>on</strong>g> use <str<strong>on</strong>g>of</str<strong>on</strong>g> biological c<strong>on</strong>trol in California may pose a risk to <str<strong>on</strong>g>the</str<strong>on</strong>g> closely related nativeSpartina foliosa, it would be an excellent opti<strong>on</strong> in o<str<strong>on</strong>g>the</str<strong>on</strong>g>r o<str<strong>on</strong>g>the</str<strong>on</strong>g>r parts <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> world where Spartina hasinvaded and where <str<strong>on</strong>g>the</str<strong>on</strong>g>re are no closely related native Spartina species. In additi<strong>on</strong> to P. marginata,o<str<strong>on</strong>g>the</str<strong>on</strong>g>r candidate bioc<strong>on</strong>trol agents from <str<strong>on</strong>g>the</str<strong>on</strong>g> Atlantic Coast are currently being investigated.Keywords: Biological c<strong>on</strong>trol, Spartina alterniflora, Spartina anglica, Prokelisia marginata,Willapa Bay, Puget SoundINTRODUCTIONClassical biological c<strong>on</strong>trol <str<strong>on</strong>g>of</str<strong>on</strong>g> a pest or weed involves<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> a natural enemy (bioc<strong>on</strong>trol agent) fromano<str<strong>on</strong>g>the</str<strong>on</strong>g>r geographic regi<strong>on</strong>. The goal is to establish apermanent populati<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> bioc<strong>on</strong>trol agent that willprovide l<strong>on</strong>g-term c<strong>on</strong>trol. During <str<strong>on</strong>g>the</str<strong>on</strong>g> last century, close to1,000 biological c<strong>on</strong>trol introducti<strong>on</strong>s for weeds were madethroughout <str<strong>on</strong>g>the</str<strong>on</strong>g> world (Julien and Griffiths 1998). Modernweed bioc<strong>on</strong>trol projects in <str<strong>on</strong>g>the</str<strong>on</strong>g> United States proceed <strong>on</strong>lyafter extensive testing <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> natural enemy followed by areview by <str<strong>on</strong>g>the</str<strong>on</strong>g> federal Technical Advisory Group <strong>on</strong>Biological C<strong>on</strong>trol <str<strong>on</strong>g>of</str<strong>on</strong>g> Weeds to ensure that it will not harmo<str<strong>on</strong>g>the</str<strong>on</strong>g>r organisms. Weed bioc<strong>on</strong>trol has proven to be a safeand <str<strong>on</strong>g>of</str<strong>on</strong>g>ten very effective method <str<strong>on</strong>g>of</str<strong>on</strong>g> l<strong>on</strong>g term c<strong>on</strong>trol <str<strong>on</strong>g>of</str<strong>on</strong>g>widespread invasive plants (Cruttwell-McFadyen 1998).Biological c<strong>on</strong>trol has both advantages anddisadvantages over traditi<strong>on</strong>al c<strong>on</strong>trol. Unlike most chemicaland mechanical approaches, biological c<strong>on</strong>trol is highlyspecific to <str<strong>on</strong>g>the</str<strong>on</strong>g> target weed. Since biological c<strong>on</strong>trol agentshave been chosen for <str<strong>on</strong>g>the</str<strong>on</strong>g>ir host specificity, <str<strong>on</strong>g>the</str<strong>on</strong>g>y will notharm o<str<strong>on</strong>g>the</str<strong>on</strong>g>r plant species intermixed with <str<strong>on</strong>g>the</str<strong>on</strong>g> weed.Biological c<strong>on</strong>trol is ec<strong>on</strong>omical over large areas. Onceestablished, <str<strong>on</strong>g>the</str<strong>on</strong>g> biological c<strong>on</strong>trol agent will reproduce,spread to new sites, and c<strong>on</strong>tinue to damage <str<strong>on</strong>g>the</str<strong>on</strong>g> plant withlittle or no additi<strong>on</strong>al input. Biological c<strong>on</strong>trols have notoxic residues or health hazards as do some herbicides.When it is successful, biological c<strong>on</strong>trol can provide apermanent soluti<strong>on</strong> to a weed problem, although it usuallymaintains a very low level <str<strong>on</strong>g>of</str<strong>on</strong>g> infestati<strong>on</strong> ra<str<strong>on</strong>g>the</str<strong>on</strong>g>r than bringingabout full eradicati<strong>on</strong>.The disadvantages <str<strong>on</strong>g>of</str<strong>on</strong>g> biological c<strong>on</strong>trol include <str<strong>on</strong>g>the</str<strong>on</strong>g>large amount <str<strong>on</strong>g>of</str<strong>on</strong>g> pre-release research that is required.Moreover, even with careful selecti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> host-specificagents, <str<strong>on</strong>g>the</str<strong>on</strong>g>re will always be some small risk to n<strong>on</strong>-targetorganisms from ei<str<strong>on</strong>g>the</str<strong>on</strong>g>r direct or indirect interacti<strong>on</strong>s.Biological c<strong>on</strong>trol is <str<strong>on</strong>g>of</str<strong>on</strong>g>ten slow in its acti<strong>on</strong>, taking severalyears and even up to a decade for an impact to be seen.Finally, <str<strong>on</strong>g>the</str<strong>on</strong>g> complexities <str<strong>on</strong>g>of</str<strong>on</strong>g> ecological interacti<strong>on</strong>s mean that<str<strong>on</strong>g>the</str<strong>on</strong>g> effectiveness <str<strong>on</strong>g>of</str<strong>on</strong>g> biological c<strong>on</strong>trol is difficult to predictahead <str<strong>on</strong>g>of</str<strong>on</strong>g> time. The probability <str<strong>on</strong>g>of</str<strong>on</strong>g> successful c<strong>on</strong>trolincreases when multiple bioc<strong>on</strong>trol agents are used (Deno<str<strong>on</strong>g>the</str<strong>on</strong>g>t al. 2003).A biological c<strong>on</strong>trol program for Spartina alternifloraand Spartina anglica in Washingt<strong>on</strong> State was developedduring <str<strong>on</strong>g>the</str<strong>on</strong>g> late 1990s. The bioc<strong>on</strong>trol agent Prokelisiamarginata, a sapsucking planthopper, was introducedbeginning in 2000. In this paper, we provide backgroundinformati<strong>on</strong> about this biological c<strong>on</strong>trol program. Then wepresent results <str<strong>on</strong>g>of</str<strong>on</strong>g> a comparis<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> performance <str<strong>on</strong>g>of</str<strong>on</strong>g> four- 267 -
Chapter 4: Spartina C<strong>on</strong>trol and Management<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> Spartinapopulati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> P. marginata imported from differentgeographic locati<strong>on</strong>s. Finally, we present possible futuredirecti<strong>on</strong>s for <str<strong>on</strong>g>the</str<strong>on</strong>g> bioc<strong>on</strong>trol program including <str<strong>on</strong>g>the</str<strong>on</strong>g> screening<str<strong>on</strong>g>of</str<strong>on</strong>g> additi<strong>on</strong>al agents from S. alterniflora’s native range.BACKGROUND ON SPARTINA BIOCONTROL INWASHINGTON STATETo date <strong>on</strong>ly <strong>on</strong>e biological c<strong>on</strong>trol agent has beenintroduced into Washingt<strong>on</strong> for c<strong>on</strong>trol <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina spp. Thedelphacid planthopper, Prokelisia marginata, wasintroduced from California into Willapa Bay beginning in2000 and into north Puget Sound in 2003. The introducti<strong>on</strong>swere made <strong>on</strong>ly after extensive testing dem<strong>on</strong>strated its highlevel <str<strong>on</strong>g>of</str<strong>on</strong>g> host specificity (Grevstad et al. 2003) and afterruling out <str<strong>on</strong>g>the</str<strong>on</strong>g> possibility that it could vector a disease (Daviset al. 2002). The project was reviewed and approved by <str<strong>on</strong>g>the</str<strong>on</strong>g>Technical Advisory Group <strong>on</strong> Biological C<strong>on</strong>trol <str<strong>on</strong>g>of</str<strong>on</strong>g> Weedsand permitted by <str<strong>on</strong>g>the</str<strong>on</strong>g> Washingt<strong>on</strong> State Department <str<strong>on</strong>g>of</str<strong>on</strong>g>Agriculture and <str<strong>on</strong>g>the</str<strong>on</strong>g> U.S. Department <str<strong>on</strong>g>of</str<strong>on</strong>g> Agriculture’sAnimal and Plant Health Inspecti<strong>on</strong> Service (USDA-APHIS).P. marginata is native to <str<strong>on</strong>g>the</str<strong>on</strong>g> Atlantic and Gulf Coasts <str<strong>on</strong>g>of</str<strong>on</strong>g>North America and also occurs in San Francisco Bay,California. Genetic analyses (R. Denno and D. Hawthorne,University <str<strong>on</strong>g>of</str<strong>on</strong>g> Maryland, pers. comm.) indicate that P.marginata was probably introduced to California from <str<strong>on</strong>g>the</str<strong>on</strong>g>East Coast in recent decades. The absence <str<strong>on</strong>g>of</str<strong>on</strong>g> Prokelisia spp.in an early 1970’s survey <str<strong>on</strong>g>of</str<strong>on</strong>g> insects <strong>on</strong> Spartina foliosa inSan Francisco Bay also supports a recent introducti<strong>on</strong> to <str<strong>on</strong>g>the</str<strong>on</strong>g>West Coast (Camer<strong>on</strong> 1972). P. marginata was selected as apromising bioc<strong>on</strong>trol agent because <str<strong>on</strong>g>of</str<strong>on</strong>g> its narrow host rangeand its known potency against S. alterniflora and S. anglica(Daehler and Str<strong>on</strong>g 1997; Wu et al. 1999). P. marginataadults and nymphs feed by sucking <str<strong>on</strong>g>the</str<strong>on</strong>g> sap from <str<strong>on</strong>g>the</str<strong>on</strong>g> plant,draining its energy supply. Spartina is also damaged by <str<strong>on</strong>g>the</str<strong>on</strong>g>scars that arise <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> leaf surface where adult females inserteggs. If high enough densities <str<strong>on</strong>g>of</str<strong>on</strong>g> P. marginata are attained,feeding and ovipositi<strong>on</strong> scars cause <str<strong>on</strong>g>the</str<strong>on</strong>g> leaves to turn brownand eventually kill <str<strong>on</strong>g>the</str<strong>on</strong>g> plant.The Spartina bioc<strong>on</strong>trol program is unique in being <str<strong>on</strong>g>the</str<strong>on</strong>g>first classical bioc<strong>on</strong>trol program to target a grass, althougho<str<strong>on</strong>g>the</str<strong>on</strong>g>rs are being c<strong>on</strong>sidered (Tewksbury et al. 2002; Wittand McC<strong>on</strong>nachie 2004). It is also <str<strong>on</strong>g>the</str<strong>on</strong>g> first applicati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>classical weed bioc<strong>on</strong>trol in a marine intertidal envir<strong>on</strong>ment.This project differs from most classical bioc<strong>on</strong>trol projectsin that <str<strong>on</strong>g>the</str<strong>on</strong>g> targeted weed is invasive in <str<strong>on</strong>g>the</str<strong>on</strong>g> same country towhich it is native (although a different regi<strong>on</strong>). Thebioc<strong>on</strong>trol agents are likewise transferred between statesra<str<strong>on</strong>g>the</str<strong>on</strong>g>r than between countries.To <str<strong>on</strong>g>the</str<strong>on</strong>g> advantage <str<strong>on</strong>g>of</str<strong>on</strong>g> a biological c<strong>on</strong>trol program,invasive Spartina in Washingt<strong>on</strong> appears to have lostresistance to herbivory since its introducti<strong>on</strong>. In greenhouseexperiments (Daehler and Str<strong>on</strong>g 1997; Wu et al. 1999;Garcia-Rossi et al. 2003), plants from <str<strong>on</strong>g>the</str<strong>on</strong>g> invasivepopulati<strong>on</strong>s in Washingt<strong>on</strong> suffer much greater biomassreducti<strong>on</strong> and mortality from P. marginata than plants fromnative locati<strong>on</strong>s. Herbivore exclusi<strong>on</strong> and additi<strong>on</strong>experiments in <str<strong>on</strong>g>the</str<strong>on</strong>g> field also dem<strong>on</strong>strate this difference inresp<strong>on</strong>se (compare Daehler and Str<strong>on</strong>g 1996 with Grevstadet al. 2003). The vulnerability <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Washingt<strong>on</strong>populati<strong>on</strong>s may be due to an evolved loss <str<strong>on</strong>g>of</str<strong>on</strong>g> resistance in<str<strong>on</strong>g>the</str<strong>on</strong>g> absence <str<strong>on</strong>g>of</str<strong>on</strong>g> herbivores (Garcia-Rossi et al. 2003). Themechanism <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina vulnerability is unknown, but it maybe related to <str<strong>on</strong>g>the</str<strong>on</strong>g> structural breakdown <str<strong>on</strong>g>of</str<strong>on</strong>g> vascular cells as aresult <str<strong>on</strong>g>of</str<strong>on</strong>g> piercing by <str<strong>on</strong>g>the</str<strong>on</strong>g> planthopper during feeding andovipositi<strong>on</strong> (Wu et al. 1999). The possibility that <str<strong>on</strong>g>the</str<strong>on</strong>g>vulnerability is due to a disease vectored by <str<strong>on</strong>g>the</str<strong>on</strong>g> planthopperwas ruled out by Davis et al. (2002).Over <str<strong>on</strong>g>the</str<strong>on</strong>g> past few years, P. marginata has been releasedat 40 locati<strong>on</strong>s in Willapa Bay and Puget Sound. Resultshave been encouraging, but not without setbacks. Followingrelease, P. marginata populati<strong>on</strong>s typically grow explosivelyduring <str<strong>on</strong>g>the</str<strong>on</strong>g>ir first summer and cause visible damage to <str<strong>on</strong>g>the</str<strong>on</strong>g>plants by fall (Grevstad et al. 2003). Local densities in somesites have exceeded 50,000 insects per m 2 . A 50% reducti<strong>on</strong>in local biomass was measured in an early field cageexperiment (Grevstad et al. 2003). A 90% reducti<strong>on</strong> in fieldseed viability was found in localized areas where P.marginata density was greater than 30 per stem (Grevstad,unpublished data). However, low survival <str<strong>on</strong>g>of</str<strong>on</strong>g> nymphs over<str<strong>on</strong>g>the</str<strong>on</strong>g> winter has prevented <str<strong>on</strong>g>the</str<strong>on</strong>g>se populati<strong>on</strong>s from building to<str<strong>on</strong>g>the</str<strong>on</strong>g> high densities over large areas required to have largescaleimpacts <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> Spartina populati<strong>on</strong>. In spite <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>summer boom, P. marginata populati<strong>on</strong>s are typically muchsmaller <str<strong>on</strong>g>the</str<strong>on</strong>g> following spring than <str<strong>on</strong>g>the</str<strong>on</strong>g>y were at <str<strong>on</strong>g>the</str<strong>on</strong>g> time <str<strong>on</strong>g>of</str<strong>on</strong>g>release. Some <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>se populati<strong>on</strong>s eventually build updensities, but many have g<strong>on</strong>e extinct and o<str<strong>on</strong>g>the</str<strong>on</strong>g>rs aredwindling or growing <strong>on</strong>ly slowly.Determining <str<strong>on</strong>g>the</str<strong>on</strong>g> best geographic source <str<strong>on</strong>g>of</str<strong>on</strong>g> P. marginataIn selecting a geographic source <str<strong>on</strong>g>of</str<strong>on</strong>g> a bioc<strong>on</strong>trol agent, itis important to c<strong>on</strong>sider <str<strong>on</strong>g>the</str<strong>on</strong>g> ways that herbivore populati<strong>on</strong>smay be locally adapted. Classical biological c<strong>on</strong>trolprograms <str<strong>on</strong>g>of</str<strong>on</strong>g>ten seek an agent source populati<strong>on</strong> from alocati<strong>on</strong> that has a climate similar to <str<strong>on</strong>g>the</str<strong>on</strong>g> regi<strong>on</strong> where it willbe introduced. However, in <str<strong>on</strong>g>the</str<strong>on</strong>g> case <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> Spartinabioc<strong>on</strong>trol program, a close match to <str<strong>on</strong>g>the</str<strong>on</strong>g> Willapa Bayclimate does not exist. The San Francisco Bay area has asimilarly moderate climate but temperatures areapproximately 5ºC warmer at all times <str<strong>on</strong>g>of</str<strong>on</strong>g> year. East Coastlocati<strong>on</strong>s have more extreme seas<strong>on</strong>ality and no locati<strong>on</strong> canmatch both winter and summer temperatures. A nor<str<strong>on</strong>g>the</str<strong>on</strong>g>asternlocati<strong>on</strong> such as Rhode Island has <str<strong>on</strong>g>the</str<strong>on</strong>g> best match during <str<strong>on</strong>g>the</str<strong>on</strong>g>summer m<strong>on</strong>ths, but a mid-Atlantic locati<strong>on</strong>, such asVirginia, has <str<strong>on</strong>g>the</str<strong>on</strong>g> best match during <str<strong>on</strong>g>the</str<strong>on</strong>g> winter.- 268 -
- Page 2 and 3:
Proceedings <stron
- Page 4 and 5:
FORWARD & ACKNOWLEDGEMENTSThe <stro
- Page 6 and 7:
TABLE OF CONTENTSForward & Acknowle
- Page 9 and 10:
Community Spartina Education and St
- Page 11 and 12:
included the docum
- Page 14:
CHAPTER ONESpartina Biology
- Page 17 and 18:
Chapter 1: Spartina Biology
- Page 19 and 20:
Chapter 1: Spartina Biology
- Page 21 and 22:
Chapter 1: Spartina Biology
- Page 23 and 24:
Chapter 1: Spartina Biology
- Page 25 and 26:
Chapter 1: Spartina Biology
- Page 28 and 29:
Proceedings <stron
- Page 30 and 31:
Proceedings <stron
- Page 32 and 33:
Proceedings <stron
- Page 34:
Proceedings <stron
- Page 37 and 38:
Chapter 1: Spartina Biology
- Page 39 and 40:
Chapter 1: Spartina Biology
- Page 42 and 43:
Proceedings <stron
- Page 44:
Proceedings <stron
- Page 47 and 48:
Chapter 1: Spartina Biology
- Page 49 and 50:
Chapter 1: Spartina Biology
- Page 51 and 52:
Chapter 1: Spartina Biology
- Page 53 and 54:
Chapter 1: Spartina Biology
- Page 55 and 56:
Chapter 1: Spartina Biology
- Page 57 and 58:
Chapter 1: Spartina Biology
- Page 60 and 61:
Proceedings <stron
- Page 62 and 63:
Proceedings <stron
- Page 64 and 65:
Proceedings <stron
- Page 66:
Proceedings <stron
- Page 69 and 70:
Chapter 1: Spartina Biology
- Page 71 and 72:
Chapter 1: Spartina Biology
- Page 74 and 75:
Proceedings <stron
- Page 76:
Proceedings <stron
- Page 79 and 80:
Chapter 2: Spartina Distribution an
- Page 81 and 82:
Chapter 2: Spartina Distribution an
- Page 83 and 84:
Chapter 2: Spartina Distribution an
- Page 86 and 87:
Proceedings <stron
- Page 88 and 89:
Proceedings <stron
- Page 90 and 91:
Proceedings <stron
- Page 92 and 93:
Proceedings <stron
- Page 94 and 95:
Proceedings <stron
- Page 96 and 97:
Proceedings <stron
- Page 98:
Proceedings <stron
- Page 101 and 102:
Chapter 2: Spartina Distribution an
- Page 103 and 104:
Chapter 2: Spartina Distribution an
- Page 105 and 106:
Chapter 2: Spartina Distribution an
- Page 108 and 109:
Proceedings <stron
- Page 110:
Proceedings <stron
- Page 113 and 114:
Chapter 2: Spartina Distribution an
- Page 115 and 116:
Chapter 2: Spartina Distribution an
- Page 117 and 118:
Chapter 2: Spartina Distribution an
- Page 119 and 120:
Chapter 2: Spartina Distribution an
- Page 122 and 123:
Proceedings <stron
- Page 124 and 125:
Proceedings <stron
- Page 126 and 127:
Proceedings <stron
- Page 128:
Proceedings <stron
- Page 131 and 132:
Chapter 2: Spartina Distribution an
- Page 134 and 135:
Proceedings <stron
- Page 136 and 137:
Proceedings <stron
- Page 138 and 139:
Proceedings <stron
- Page 140:
CHAPTER THREEEcosystem Effects <str
- Page 143 and 144:
Chapter 3: Ecosystem Effects <stron
- Page 145 and 146:
Chapter 3: Ecosystem Effects <stron
- Page 148 and 149:
Proceedings <stron
- Page 150 and 151:
Proceedings <stron
- Page 152:
Proceedings <stron
- Page 155 and 156:
Chapter 3: Ecosystem Effects <stron
- Page 157 and 158:
Chapter 3: Ecosystem Effects <stron
- Page 160 and 161:
Proceedings <stron
- Page 162 and 163:
Proceedings <stron
- Page 164:
Proceedings <stron
- Page 167 and 168:
Chapter 3: Ecosystem Effects <stron
- Page 169 and 170:
Chapter 3: Ecosystem Effects <stron
- Page 171 and 172:
Chapter 3: Ecosystem Effects <stron
- Page 174 and 175:
Proceedings <stron
- Page 176:
Proceedings <stron
- Page 179 and 180:
Chapter 3: Ecosystem Effects <stron
- Page 181 and 182:
Chapter 3: Ecosystem Effects <stron
- Page 184 and 185:
Proceedings <stron
- Page 186 and 187:
Proceedings <stron
- Page 188 and 189:
Proceedings <stron
- Page 190 and 191:
Proceedings <stron
- Page 192 and 193:
Proceedings <stron
- Page 194 and 195:
Proceedings <stron
- Page 196:
Proceedings <stron
- Page 199 and 200:
Chapter 3: Ecosystem Effects <stron
- Page 201 and 202:
Chapter 3: Ecosystem Effects <stron
- Page 204 and 205:
Proceedings <stron
- Page 206 and 207:
Proceedings <stron
- Page 208 and 209:
Proceedings <stron
- Page 210 and 211:
Proceedings <stron
- Page 212:
Proceedings <stron
- Page 216 and 217:
Proceedings <stron
- Page 218 and 219:
Proceedings <stron
- Page 220 and 221:
Proceedings <stron
- Page 222 and 223:
Proceedings <stron
- Page 224 and 225:
Proceedings <stron
- Page 226 and 227:
Proceedings <stron
- Page 228 and 229:
Proceedings <stron
- Page 230 and 231: Proceedings <stron
- Page 232 and 233: Proceedings <stron
- Page 234 and 235: Proceedings <stron
- Page 236 and 237: Proceedings <stron
- Page 238 and 239: Proceedings <stron
- Page 240 and 241: Proceedings <stron
- Page 242 and 243: Proceedings <stron
- Page 244 and 245: Proceedings <stron
- Page 246: Proceedings <stron
- Page 249 and 250: Chapter 4: Spartina Control and Man
- Page 251 and 252: Chapter 4: Spartina Control and Man
- Page 253 and 254: Chapter 4: Spartina Control and Man
- Page 255 and 256: Chapter 4: Spartina Control and Man
- Page 257 and 258: Chapter 4: Spartina Control and Man
- Page 259 and 260: Chapter 4: Spartina Control and Man
- Page 261 and 262: Chapter 4: Spartina Control and Man
- Page 263 and 264: Chapter 4: Spartina Control and Man
- Page 265 and 266: Chapter 4: Spartina Control and Man
- Page 267 and 268: Chapter 4: Spartina Control and Man
- Page 269 and 270: Chapter 4: Spartina Control and Man
- Page 271 and 272: Chapter 4: Spartina Control and Man
- Page 273 and 274: Chapter 4: Spartina Control and Man
- Page 276 and 277: Proceedings <stron
- Page 278 and 279: Proceedings <stron
- Page 282 and 283: Proceedings <stron
- Page 284 and 285: Proceedings <stron
- Page 286 and 287: Proceedings <stron
- Page 288 and 289: Proceedings <stron
- Page 290: Proceedings <stron