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Supporting : Interdisciplinary perspective shows Spartina as a distinct solid Alejandro Bortolus, Paul Adam, Janine Adams, Malika Ainouche, Debra Ayres, Mark Bertness, Tjeerd Bouma, John Bruno, Isabel Caçador, James Carlton, et al.

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Alejandro Bortolus, Paul Adam, Janine Adams, Malika Ainouche, Debra Ayres, et al.. Supporting Spartina : Interdisciplinary perspective shows Spartina as a distinct solid genus. , Ecological Society of America, 2019, 100 (11), pp.e02863. ￿10.1002/ecy.2863￿. ￿hal-02355287￿

HAL Id: hal-02355287 https://hal-univ-rennes1.archives-ouvertes.fr/hal-02355287 Submitted on 18 Nov 2019

HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Hughes Bertness Mark D. Alejandro Bortolus Thi 3 2 (U9120ACD), Chubut, Madryn Ecosistemas Continentales (IPEEC),Blvd. Brown2915,Puerto CONICET, 1 Sytsma Muñoz-Rodríguez Figueroa M. Castillo Running Supporting Head: Spartina Sydney, New South Wales, Australia Sydney, Australia SouthWales, New S. Weis Salmon Pennings

Department of Botany, Nels Department ofBotany, andEnvironmental School ofBiological, Earth Supporting Accepted Co en Ambientes Grupo deEcología Article 33 19 4 37 , Evangelina Schwindt 11 , Stijn Temmerman 26 , EnriqueMateos-Naranjo

, Joel Gerwein , Joel , AndreaPickart 11 , Cesar S.B. Costa , CesarS.B. Spartina 6 22 , Tjeerd J. Bouma J. , Tjeerd 1,38 , Francisco J.J. Nieva J.J. , Francisco , Paul Adam : Interdisciplinary perspective shows shows perspective : Interdisciplinary 15 , Alan J. Gray , Alan J. 27 34 , SusanaRedondo-Gómez on Mandela University, Port Elizabeth, South PortElizabeth, on MandelaUniversity, , R.Eugene Turner 29 12 , Brian R. Silliman , Anthony J. Davy , Anthony J. 2 11 , Janine B. Adams , Janine

7 , IrvingA. Mendelssohn , John F., Bruno John steros (GEAC), Instituto steros (GEAC), Patag 16 22 , Edwin D.Grosholz , Edwin , Lisa A.Levin Science, University of New South Wales, SouthWales, ofNew University Science, 20 , IvanValiela 13 30 , Linda Deegan 8 , ErikE.Sotka , Isabel Caçador 3 , Malika L., Malika Ainouche 11 , DavidM.Richardson 23 , Bo Li, Bo 20 Spartina 17 , T.Morris James 35 , Sally D. Hacker , SallyD. , Michael P. Weinstein P. , Michael 24 31 14 , Wenwen Liu, Wenwen ónico para elEstudiodelos ónico , Clive Stace , Clive 9 , Bernardo Duarte , James T.Carlton, James asa distinct solidgenus 4 , DebraAyres 28 21 , Armel 18 32 , AdolfoF. 25 , A.Randall , Mark , Steven C. C. , Steven 9 , Enrique 10 36 , Jesus , Jesus , Judith , Judith 5 , USA USA LA, USA 20 19 18 17 16 15 14 13 12 11 10 Lisbon, Portugal Lisbon, CampoGrande, 9 8 Netherlands The Utrecht University, Geography, Department ofPhysical Geosciences, Faculty of and TheNetherlands; 140,4400ACYerseke, P.O.Box University, Utrecht (NIOZ), 7 6 5 4 Campus de Beaulieu, 35 042 Campus deBeaulieu, 35 . MARE - Marine and Environmental MARE -MarineandEnvironmental DepartmentUniversity ofBiology, Chapel NorthCarolina, Hill,NC,USA of Department ofEstuarine and Delta Netherlands systems, Royal Institute Sea for Research and E Department ofEcology Evolution andEcology,University UMRCNRSBiodiversity 6553Ecosystems, Evol School ofBiological Sciences,Univer inburgh Research Station,Scotland, UK Hydrology, Research Edinburgh and Centre forEcology Coastal State California Center, 129W HoleResearch Woods Accepted Rouge, Baton University, Louisiana State Sciences, andCoastal ofOceanography Department Department ofMarine Integrative Biology, Oregon Stat Department of Department ofEnvironmental Science Policy, and University Ri Universidade Federal do Departamento de BiologíayUniversidad Vegetal Ecología, de Sevilla, Spain Sevilla, Maritime Studies Program, Williams College Article tern University, Nahant, MA, USA MA,USA Nahant, University, tern Northeas and EnvironmentalScience, Conservancy, CA, USA Conservancy, CA,USA o Grande-FURG, Campus Carre Campus o Grande-FURG, Rennes Cedex, France France Rennes Cedex, volutionary Biology, Brown University, Providence, RI, USA USA RI, Brownvolutionary Biology, Providence, University, of California, Davis, CA,USA California, of Sciences Centre, Faculty Sc Sciences Centre, of oods Hole Road, Falmouth, MA, USA oods HoleRoad,Falmouth, MA,USA sity of East Anglia, Norwich, UK UK Anglia, Norwich, sity ofEast Mystic Seaport, Mystic,CT, USA e University, 3029Cordley Corvallis,OR, Hall, ution (ECOBIO), University of Rennes 1, ution (ECOBIO), ofRennes University iros, Rio Grande, RS,Brazil iros, RioGrande, of California, Davis, CA, USA Davis,CA,USA California, of iences of the University of of the University iences Argentina USA Arcata, CA, 33 32 SC, USA 31 USA NC, Beaufort, University, 30 29 28 27 26 Fujian 361102, China University, Xiamen the EnvironmentandEcology, 25 200438, China Institute of Science and Coastal Ecosystems Research Station ofthe Yangtze River ,Institute ofBiodiversity 24 InstitutionScripps La ofOceanography, CA,USA Jolla, 23 22 Columbia, SC,USA 21 Matieland, South Africa Matieland, SouthAfrica . Portland State University, Portland, OR, USA Portland StateUniversity, Portland,OR,USA Bo of Invasion Department Biology, Centre for Grice Marine Laboratory and the Department of Department Laboratoryand the Grice Marine Duke Nicholas School oftheEnvironment, and Conservation, Division ofMarineScience Instituto de Biología deOrganismos Ma U.S. Fish andWildlife Service, Humboldt Department ofBiologyand MinistryKey ofthe ofEducation Laboratory Ministry of EducationKey Laboratory Biodiv for DivisionandCenter Integrative Oceanography Integradas, de Ciencias Departamento BelleInstitute Baruch forMarine Coastal and Sciences, Carolina, University ofSouth AcceptedIP14Lane, Sufolk Larters UK 5HB, MiddlewoodGreen, Apletree House, Article Eco-Chongming, School of Life Eco-Chongming,of Sc School Biochemistry, University of Houston, Houston, TX77204,USA Biochemistry, ofHouston, University Universidad de Huelva, Huelva, Spain Huelva,Spain deHuelva, Universidad rinos (IBIOMAR-CONICET), Madryn, Puerto Bay National Wildlife Refuge, 6800 Lanphere Refuge, 6800 Rd., National Wildlife Bay for Coastalfor and Wetland Ec for Marine and Conservation, Biodiversity and Marine for tany and Zoology, Stellenbosch University, University, Zoology,tany Stellenbosch and Biology, Charleston, College Charleston, of ersity Science and Ecological Engineering, Engineering, andEcological ersity Science iences, Fudan University, Shanghai University, Shanghai iences, Fudan osystems, Collegeof genus asthe betreated should simplyefficiently and considerth We interdisciplinary perspective. this debatewewillencourage pos phylogenetic insights ofthe subjective and form proposal tochange Spartina Spartina history, economics, and sociology. There isnorationale socompelling the tosubsume name biologi ecology, experimental multiple disciplines, including ecology,evolutionary biology, molecular biology, biogeography, Spartina genera included inthe previously others) Sporobolinae confir study phylogenetic Ina DNA-based 2014 ABSTRACT 38 37 36 35 34 . Corresponding Author E- Corresponding Author Accepted ArticleDepartment ofBiological Cons New Sciences Jersey Marine Biolog Marine The Ecosystems Center, Un ResearchGroup, Ecosystem Management for over 200 years. We do not agree with the subjective arguments underlying the years. withthesubjectivearguments donotagree We forover200 asasubgenusthatcouldrivalthestriki species have contributed substantially (and specieshavecontributed proposed the creation of a large monophyletic genus monophyletic ofalarge proposedthecreation to Spartina mail: [email protected] mail: [email protected] Sciences, Rutgers University, Newark, NJ, USA NJ, Newark, University, Sciences,Rutgers cal invasions, environmental ma cal invasions,environmental We understand the importance ofboththeobjective understandSporobolus. We theimportance itive feedbackthatwillstrengthen ortium, Sandy Hook, New Jersey, USA USA Hook,NewJersey, ortium, Sandy ical Laboratory, Woods Hole, MA, USA Laboratory, Hole,MA, Woods ical at the stronglyclade distinct,monophyleticat the iversity of Antwerp, Antwerp, Belgium Antwerp, iversity ofAntwerp, ming the paraphyly of the grass subtribe ofthe theparaphyly ming alized nomenclature and hope that by opening thatby opening nomenclatureandhope alized useofthename and ng, global iconichistory Spartina continue contributing) to our knowledge in toourknowledge continue contributing) Spartina , . nagement, ecology, restoration taxonomic decisionswithan taxonomic , and Sporobolus. , including (among Spartina

genera. Their proposal was accepted by a acceptedby was Their proposal genera. conserving proposed they nomenclatural priority, of retain the name priority Spartina genusshould ICBN Thenewly combinedmonophyletic the rules. Grasses 113.1890)under 169.1810)and (Sporobolus R.Brown, Prodr. Spartina The name precedence. forhistorical Linnaeus) by Plantarum ofSpecies publication (first guiding inthisnomenclature is principle al. 2018).The Inte the and detailedin namingThe are taxa rulesInternational for by governed the forPlant Association withabasicchromoso dodecaploid species ofthispolyploid the emergence understanding of the closest relative(sister) lineages of morphological, andchromosomenumb physiological that provides critical insights intotheevolutionaryhistory ofthe (Peterson etal.2014 genera included inthe previously others) species Sporobolinae confir study phylogenetic Ina DNA-based 2014 decisions, integrative analysis words: Key .

AcceptedSporobolus Article (Spartina Schreb., Gen. Pl. ed. 8[a]. Gen.Pl.ed.8[a]. (SpartinaSchreb., cordgrass, , coastal ecology, botanical nomenclature, interdisciplinary interdisciplinary nomenclature, botanical ecology, coastal marsh, salt cordgrass, proposed the creation of a large monophyletic genus Sporobolus monophyletic ofalarge proposedthecreation ) recognized that the name name the that Peterson etal.(2014a)recognized in1810.While search is an important contribution isanimportant research Thiscomprehensivephylogenetic a,b). first published in 1789, that predates recognition anddescription firstpublishedin 1789, thatpredatesrecognition rnational Code of Botanical No rnational Code Spartina Permanent Nomenclature Committeeofthe Nomenclature Permanent me number of x=10 (Ainouche et al. 2009). et al.2009). (Ainouche me numberofx=10 43. over 1789) hashistoricalpriority Calamovilfa clade which includes only tetraploid to tetraploid whichincludesonly clade ming the paraphyly of the grass subtribe ofthe theparaphyly ming Spartina , which isofcrucial importance for Sporobolus er evolution. It also priority , Calamovilfa (Calamovilfa, A. Gray), Hack.,True against against , with a formal base date of 1753 basedateof , withaformal Sporobolinae menclature (ICBN; Turland et Turland (ICBN; menclature aswellother Spartina , and Sporobolus provides a clear picture picture provides aclear , encompassing its its , encompassing , including (among Spartina Sporobolus had

andsociology. history,ecology, economics, environmen ecology, experimental biogeography, biology,knowledge inmultipledisciplines,including molecular biology, evolutionary contributed substantia genus 2008) andthe has Indian Oceans. Some for Antarcticaare present and remote onsome ofthe islands most ofthe Pacific Atlantic, and Peterson etal.2014 et al.2007, Fortune etal.2002, (Baumel clade amonophyletic representing (Mobberley 1956), The genus multidisciplinary often atefforts, immense cost. legacy significant of will servetounderminethe proposal tochange the rationale underlying (M changes” “disadvantageous nomenclatural Spartina and well-rooted scientific, history of cultural andrelevance socio-economic implicit inthe name overlooking thelong tin etal.2015),butalso MYA;Rousseau-Gueu sister cladesometimes12-20 (e.g. clades divergent highly encompasses genus that large extremely ofan creation including the notonly presents majordisadvantages This proposal,however, name Sporobolus Th forPlantTaxonomy.International Association . Accepted Article Spartina . The important criterion for rejection or conservation ofbotan or rejection . Theimportantcriterionfor Spartina would avoid taxonomic destabilization, wouldavoidtaxonomic notin would necessitate many new binomialco new wouldnecessitatemany Schreb. is a morphologically we isamorphologically Schreb. ). Its species are abundant and widely and abundant speciesare Its a). species are among the most studied plant species (e.g., Pyšek et al. al. et Pyšek species (e.g., plant studied most the among are species Spartina to Spartina acNeill etal.2015).Wedonotagree withthe acNeill lly (and itcontinuescontributing)lly toour advances achieved after decades of worldwide ofworldwide decades after achieved advances tal management, restoration ecology, invasion ecology, management, restoration tal e authors argued that conserving the name the thatconserving e authorsargued ll-circumscribed grll-circumscribed Sporobolus mbinations and heterotypic synonyms. synonyms. mbinations andheterotypic g that shiftingall names tothe senior distributed on every continent except continentexcept distributedonevery . We argue that this change change this argue that . We divergedfromits Spartina ical namesistoavoidof ical oup ofgrasses oup through trophic interactions includingthrough thenursery trophicinteractions the global onoutwelling-pulsing paradigm eventsin integrative ecology, “connectivity” the biodegradation pollutants, oforganic the negative vs.positive interactions incommunityecology, The smoothcordgrass 1993, Bertness 1999, Ganetal.2009,Silliman al. 2009). producti primary andsecondary tocoastalaccreti relative conservation ecology mirages the“ecological inspiring engineering, Spartina coastal bi 1908), aswellinpioneerworkson 1821,Barlett (Eliot colonial figuredin the andbotany inagriculture prominently inbiogeomorphology well asadvances 2005,Hacker andHacker biology (Dethier theroleof inourunderstanding of keyadvances Silliman andBertness Silliman 2002, ecology inecosystem debate up" processes envir of 2003), thefunctioning andKr 1981,Weinstein 1979, Mendelssohnetal. debatesonthedy zonation, in inmodernecology, concepts andperspectives spec iconic This . and subtropical temperate . Accepted Article species have played a major role inl a majorrole specieshaveplayed Spartina alterniflora Spartina namics of soilnutrientsinmarine namics of onmental eutrophication and the paramount "top-down andtheparamount onmental eutrophication vs. bottom- on of fauna species worldwide (Costa and Davy 1992,Adam Davy worldwide(Costa and species on offauna and Bortolus 2003,Valiela2015). (Morris et al. 2002). Species such as as al. 2002).Speciessuch et (Morris and Dethier 2006,2009,RilovandCrooks2009)as and Dethier , for instance, is a well-known salt marshplanton , forinstance, salt isawell-known (Bertness et al. 2001, Mendelssohn andMorris2001, etal.2001,Mendelssohn (Bertness ong-term studies of coastal environmental environmental studies ofcoastal ong-term hypothesis” (Bortolus etal.2015), andin (Bortolus hypothesis” on and marsh creation, as well asoflinkages aswell to on andmarshcreation, ogeography (Chapman 1960). In moderntimes ogeography (Chapman 1960). cluding studies on the causal processes of plant processesof studiesonthecausal cluding biotic resistance and legacy effects ininvasion effects and legacy resistance biotic eeger 2000, Beck et al. 2001, Bruno et al. al. 2001,Bruno et Beck et eeger 2000, role hypothesis role (Haine ies has inspired mostinsightful some ofthe and terrestrial ecosystems, the ecosystems, andterrestrial It has further fostered s 1979, Valiela and Teal ValielaandTeal s 1979, S. alterniflora

, South American Atlantic coast,China, coast, SouthAmericanAtlantic AmericanPacific North inEurope,the species aggressive non-native asan is widely regarded speciesisoneofth etal.2018).This Schwarz etal.2007, al.2005,2013,Temmerman et (Bouma organism traitsinlandscapeformation interdisciplinary field of “BioGeoMorphology”, oftheergot fungus thelong-termepidemic toreconstruct theevidence 2013). S.anglicaprovided etal. al. 2004,2012,Kueffer andduplicationinplants(Ainoucheet genome merger recurrent hybridization and makes thislineage following event recentspeciation This Ayres 2013). and Ainouche etal.2009,Strong native European genomeF1 hybrid fromofthe doubling al. 2004).Thisspeciesderived ofre anglica, knownascommoncordgrass, isamodel of Other species (Bertness 1991), microevolution ofclonalpl (Bertness 1991),microevolution work foundational including in marineecology, cordgrass The saltmeadow al. 2015). Ayres(Li et 2009,Wan and et al.2009,Strong impacts in striking, socio-economic cascading These invasionsresulted secondary productivity. ecosystems vast coastal-marine Asia (Qiu2013)andthe in recorded events ever alterniflora . Accepted Article and are responsible for some ofthe forsome S. anglicaareresponsible Spartina and the introduced American American introduced S. maritimaandthe idered a key model speciesinthe Ita key model etal.1998). isalsoconsidered (Raybould have also made critical contri havealsomade S. patens and alteredcoastal geomorphology has long been central to a number of important experiments experiments ofimportant central toa number haslong been ants (Silander 1979) and the physiological ants (Silander1979)and thephysiological New Zealand, Australia and South Africa. In fact, In fact, Africa. andSouth Zealand, Australia New e “100 World's Worst Invasiveand AlienSpecies” World's Worst “100 e al. 2009, Saarela 2012, Luqu 2009, Saarela2012, al. designed to understand coastal plant zonation plantzonation coastal designed tounderstand demonstrating the crucial importanceof crucial demonstrating the Americas (Bortolus et al. 2015) having reshaped reshaped etal.2015)having Americas (Bortolus a model system for studying the effects of effects modelsystem the a forstudying cent allopolyploid speciation (Ainouche et (Ainouche allopolyploidcent speciation butions toscie largest continental-scale bioinvasion continental-scale largest S. alterniflora , biodiversity, and primary and and primary , biodiversity, ntific knowledge. S. xtownsendii (Gray etal.1991, (Gray e etal.,2014,Yin ofthe Spartina S. Spartina participa “Spartina Watchers” scientists. The USAPacific coast,instance,commonly for is patrolled by volunteers called environmentalists.In itsintroduced range, itis the general public wholive along coastlinesIn the world. range itsnative around itisbeloved by Importantly, Spartina were achieved by studying by achieved were to soilanoxia,drainage,and al. 2008,Castilloet2010,2018).Substa Fortune et atunpr spreading aggressively before decades may fallowfor lay that introducedspecies showing andSpain USA oceans, withworkinthe its hybridseluc alsopermitted environment new typeofcoastal akey dominantina S.densiflora isdescribedas Theaustralcordgrass Sueiro etal.2012,2013). Pacific andtheSouthwes theNorth along andintroducedinvertebrates native for conditions andtheconsequences physical with etal. andSage,2015,Friesen (Carpita remediation projects toidentify st andinstudiesdirected research, genot ondifferent in predictivemodelsfocusing prairie cordgrass Morris 1984,Pennings Bertness, 2001).The and levels toxic reach can and thelackofoxygen mechanisms forthe intra-and extra-cellular regulation inhabitats where hypersaline conditions . Accepted Article S. foliosa populations.This hasbeenrepeated and S.densiflora is one of a small handful of speciesth isoneofasmallhandful tern Atlantic respectively (Levin tern Atlanticrespectively Spartina ting inearly-detection activ idation of the complex routes non-nativespeciesfollowacross routes idation ofthecomplex illustrate the major control that these species can exert on soil chemistry, and how these processes influence plant growth, plant influence howtheseprocesses soil chemistry, and named “rocky marshes” (Bortolu marshes” named “rocky species (Weinstein and Kreeger 2001). 2001). andKreeger (Weinstein species ly acknowledged at the “International “International atthe acknowledged ly for most species (Burdick and Mendelssohn1987, and most species(Burdick for ress tolerance genes,ress toleranceas well asinpollutant ecedented rates (Borto rates ecedented the focus ofenvironmental action by citizen ypes inbioenergy biomassproduction forwoody ntial advances in understanding adaptations adaptations ntial advancesinunderstanding ities to discover new non-native ities todiscover et al. 2006, Whitcraft and Levinand 2007, etal.2006,Whitcraft 2015). Manipulative experiments experiments 2015). Manipulative at is known by itssc at isknownby s etal.2009).Thisspeciesand is currently used iscurrently used S. pectinata lus 2006,Ayres etal.2008, ientific nameto Spartina

of broad consensus, changing changing of broadconsensus, Sporobolus adopted (https://www.tropicos.org/) unilaterally have Ge databasessuchas While theimpactson global distribution, increasing todiscussthe countries since 1990indifferent environmenta professionalscientistsand attended by Conference”, et al. 2019, Infante-Izquierdo etal.2019) Infante-Izquierdo et al.2019, Sporobolus international scientific community. Clearly,thisisnotthe case withthe change of The ultimatetestofthevalue all th required tocite arethis was a cited review if article, inthisreport, the pagesan entire of issue journal wouldbe papers have perspecies written been on used undertheTitlecategory: 1,507for Spartina 200 inSporobolus Giventhenum asaTopic. “Sporobolus” found using were while690papers Topic “Spartina”, underthe 2019) showed4,626papers Spartina alterniflora using listingsshows no(zero) alterniflora " 7,290usesof records 16,2019) accessedMay (https://scholar.google.com/, scientific intheprimary peer-reviewed decade . Accepted Article than . As one final example, whilethiswork . Asonefinalexample, " versus 11 (eleven) uses of " " usesof versus11(eleven) Sporobolus (Peterson et al 2014a) this works out (Petersonetal2014a)thisworks has search a 189 listings. Moreover, e literature producedon e literature . The discrepancy becomes more pronounced when the search termis thesearch when morepronounced becomes . Thediscrepancy nBank (https://www.ncbi.nlm.ni of a taxonomic revision isits revision of ataxonomic Spartina to Spartina Spartina ber of species in each genus, 17in ineach ber ofspecies Sporobolus alterniflorus Sporobolus that include taxonomic updates on includetaxonomic that Sporobolus since the genus was first described in1789. sincethegenuswas Spartina human society, and possible solutions. andpossiblesolutions. human society, literature. On the cont Onthe literature. problems associatedwiththisplantgroup, its vs184for than was under review, two papers (Gallego- Tévar Tévar (Gallego- review, twopapers was under has not found support for nearly half a hasnotfoundsupportfornearly half Sporobolus inthetitlebetween2015and2019,while to 8x more papers per species on perspecies more papers to8x in Web of Science (accessed May 20, May (accessed ofScience in Web acceptance and application by the application by the and acceptance h.gov/genbank/) andTropicos h.gov/genbank/) Sporobolus , givingan illusory impression " ofScience since2015.Web . While only a few examples only examples afew . While l managers, and held regularly andheldregularly l managers, rary, Googlerary, Scholar – almost 100x more –almost100x Spartina Spartina Spartina Spartina andabout species species for with otherexperts. Peterson etal.(2014 Within anthropology. literature and anddisciplines,including history, a striking ofstakeholders biology range asacross aswell name name tosubsumethe no rationalesocompelling committee rulesonthisquestion.Thereisclearly genus asa orsu isused to whetheraname mol relationships derivedfrom not genus does subgenus as a employed or Sporobolinae new subgeneric classification..." retaining (2014b) proposed Spartina nomen while potentially further any restraining Spartinafor replacing of theacceptance toinduce likely are websites) databases It isofnosmallcon 2019). (Stace the case Isles” British ofthe Flora the “New werereviewedpublished by and peer respected focused systematics. journals onplant Similarly, . Accepted Article as a subgenus thatcouldrivalthestri Spartinaasasubgenus Spartina is a deeply-nested, strongly supported clade (Peterson etal.2014 (Peterson clade strongly supported isadeeply-nested, and subgeneric classification of classification andsubgeneric rings as a signature name in ecology and name inecology rings asasignature years.name Spartina The forover200 ) supply good basis for (re)naming other (re)naming good basisfor b) supply Spartina ecular-based (ormorphological-bas ecular-based oposed classification ofthe classification "A proposed (titleofpaper)and this context, the thiscontext, phylogene after retains thenameSpartina considering publishedin2019 as a subgenus of asasubgenus bgenus are subjective; nocommission,congress,are subjective; or bgenus cern that globally popular (digital open access openaccess globally popular(digital cern that change any conclusions aboutevolutionary any change Sporobolus and sound open debate. discussionsandsoundopendebate. clatural king, global iconic history and use of the and useofthe globalking, iconichistory Sporobolus " (Table 2 legend)). Whether a name is aname Whether " (Table2legend)). Sporobolinae tic relationships providedbytic relationships ("A molecular phylogeny and phylogeny ("Amolecular ed) phylogenies. Decisionsas phylogenies. ed) a,b). Petersonetal. clades in agreement agreement in clades Sporobolus , the Editor-in-Chief who suppliedthe excellent Editor-in-Chief who insightwork. thattruly improvedour which they are affiliated.We are alsograteful deeply totwo anonymous reviewerswell as to theorgani reflect personal, anddonotnecessarily improving ourreasoningand encouraging ustoinitiate thisdebate. positionsare The authors’ forsh UK andUruguay Spain, Argentina, USA, stude colleagues, many the to grateful are We Acknowledgments genus asthe treated be efficiently et al.2014a,shouldsimply and assubgenus designated clade well-knit distinct the that consider therefore, should continuetolinkscientists, citizensandenvironmentalmanagers impacts.Thename global 2019)withprofound (IPBES change innature marshes globally as well as contri Spartina impactinsomany suchanimmense ever provided d strengthen taxonomic positive feedbackthatwill will encourage we thisdebate opening ourhopeisthatbyscientific names arehypotheses, such thatmayaccompany nomenclature formalized understandtheim indeed, wefully ge as a should notbeinterpreted but,ra nomenclatural viewpoints criti deserves question, whichwefeel a on insights. said,onthisreportwehavefocused phylogenetic This thevalueofmolecula recognize unequivocallyWe . Accepted Article , intimately tothe linked initial recognition of neral argument for thesystematic for argument neral ther, the contrary (see Bortolus 2008, 2012). Thus,ourwork Bortolus 2008,2012). (see ther, thecontrary buting to our understanding ofth buting toourunderstanding portance of both phylogenetic re phylogenetic portance ofboth cal attention. lnthissense,we cal attention. nts and eight anonymous expert taxonomists from taxonomists eightnts and expert anonymous aring their opinions, perspectives andideas, aring theiropinions,perspectives zations or networks they represent or with or ornetworksthey represent zations ecisions. Few taxa, inbotany zoology,ecisions. Few or have taxa, scientificandnon-scie r work that may provide thebasisforobjective r workthatmay provide advances (Thomson et al. 2018).Giventhat (Thomsonet advances the ecological importance coastal of salt rejection of name changes — ofnamechanges rejection Spartina e top-five direct drivers of directdrivers e top-five solutions and of the resulting andoftheresulting solutions donotseektostiflediffering subjective nomenclatural nomenclatural subjective across the globe. across We, Spartina ntific fieldsashas ntific . Spartina haslinkedand by Peterson Ainouche, M., H. Chelaifa, J. Ferreira DeCarv J. Ainouche, M.,H.Chelaifa, Baumel, A., M. Ainouche, R. J. Bayer, A-K. Ainouche, and M-T. Misset.2002.Molecular andM-T. Bayer, Ainouche, R.J. A-K. Baumel, A.,M.Ainouche, grass in 1908.Theuseof “wildrice” A.W. Barlett, Blum, C.M.Sloop, M. P. Ayres, K.Zaremba, J. Bailey, D.R.,E.K.Grotkopp, J. C.K.Anttila, L., A. Salmon,C.Pari Ainouche, M. P.Fortune, Baumel, andA.Salmon.2004.Biolog Ainouche, M.,A. Studies Cambridge Adam, P.1993.Saltmarshecology. Cited Literature .

AcceptedEvolution 22:303–3014. Phylogenetics and genus species fromthe Phylogeny ofhybridizing BritishGuiana1:5–10. of Agriculture of 713–719 US nativeBay, S.foliosainSanFrancisco California, and D.R.Strong.Hybr 2008. USA. New York, 225–244 in 2012. Polyploid in evolution Spartina ArticleRicou, andM.-T.Misset.2009.Hybridizati 484. Bi genomes. allopolyploid changes thataffect genomics: Cambridge. () Biological Invasions 11:1159–1173. Biological (Poaceae) Schreb. a natural model system for analyzing early evolutionary early analyzing system for model Spartina anglicaSchreb.anatural P.S.Soltis and Springer, D.E.Soltis,editors.Polyploidy Evolution. and Genome Spartina idization betweeninvasive idization : Dealing with highly genomes. Pages: Dealingredundant hybrid with on, polyploidyLessons from andinvasion: alho, S.Bellot, andA.Salmon. alho, A.Ainouche, sod, M. A. Grandbastien, K. Fukunaga, K.M. K. Fukunaga, sod, M.A.Grandbastien, ological Journal of the Linnean 82:475– Society ofthe ological Journal preventing coast erosion. Journal oftheBoard Journal erosion. coast preventing Spartina in Ecology. Cambridge University Press, University in Ecology. Cambridge ical relevance of polyploidy, ecology to ecology ofpolyploidy, relevance ical A. American Journal ofBotany 95: A. AmericanJournal (Poaceae) and (Poaceae) Schreb. (Poaceae). Molecular Molecular (Poaceae). Schreb. Bouma, T. J., M. B. De Vries, E. Low, G. Peralta, I. C. Tánczos, J. van de Koppel, and P. M. J. van I.de Koppel, andP.M.J. Low, J. Vries, E. G.Peralta, M.B.Bouma, C.Tánczos, De T.J., coasts: T.Carlton,andE.Schwindt.2015.ReimaginingBortolus, SouthAmerican A.,J. of 2009.Acharacterization L.Idaszkin. Bouza, andY. P. Bortolus,J. A.,E.Schwindt, us reliably Bortolus, authorsto A.2012.Guiding sciences:theunwanted inthebiological cascades Bortolus, A.2008.Error cordgrass austral Bortolus, A.2006.The 2001.MarineCommunity editors. Ecology. Bertness, andM.E.Hay, M.D.,S.D.Gaines, ofAtlantic Ecology Bertness, M.D.1999.The Zonation of Bertness, M.D.1991. K. Beck,L.W. Able,L. JR., D. Childers, M.W.,K. Heck, D.B. B. Eggleston, B. M.Gillanders, .

Accepted Ecology 86:2187–2199. macrophytes. emerging relatedtoecosystem Herman. 2005.Trade-offs Distributions 21:1267–1283. Unveiling the hidden history invasion an iconic of ecological engineer.and Diversity 29:772–780. Wetlands Patagonian marshes. salt Evolution 27:418. Ambio37:114–118. ecology. bad taxonomy in and naturalhistory. Journa ArticleSinauer Associates,Sunder Massachusetts, USA. 72:138–148. salt marsh.Ecology 51:633–641. Bioscience fishandinvertebrates. nurseries for 2001.Theidentification,conservation, Weinstein. Minell L.Hoshino, T.J. Halpern, C.G.Hays, l of Biogeography 33:158–68. 33:158–68. l ofBiogeography land, Massachusetts, USA. USA. land, Massachusetts, Spartina densifloraBrong.: itstaxonomy, biogeography shorelines. Sinauer Associates, Sunderland, SinauerAssociates, shorelines. o, Orth,P.F.R. Sheridan,andM.P. J. e taxonomic names.Tr taxonomic e engineering: A case study on stiffnessof study on Acase engineering: and and management of estuarine and marine andmarine ofestuarine and management Spartina alterniflora ends inEcology and consequences of using ofusing consequences in a New England inaNewEngland Carpita, N.andR.F.2015.Plantsbi Sage. Chapman, V. J. 1960. Salt marshes and salt deserts of the world. Interscience Pub. Inc., Interscience Pub. 1960.Salt New oftheworld. marshesChapman, V.J. andsaltdeserts Cast M.,S.Dréano,J. Castillo, J. Castillo, M.,D.R.Ayres,J. Leira-Doce, P. Bailey, J. Blum, D.R.Strong,Luque, M. and T. E. I.Mendelssohn. 1987.Wate and A. Burdick, D.M. Stachowicz,andM.D.Bert F.,Bruno,J. J. J. L. S.Temmerman, A. vanDu Bouma, T.J., .

Accepted York, USA. 5007. Paci populations invading the diversity contrasts withhigh phenotypic variabilityin heptaploid Lima, Keller,O. S.Dréano, Rousseau, J. Distributions 16:547–558. Spartina densifloraandnative ofhybrids with 2010.Theproduction Figueroa. 66:4093–4095. Articlepopulations ofSpartina 18:119–125. theory.and Evolution Trends inEcology 180–181:57–65. Geomorphology Aflum feedbacks: dependent bio-geomorphic Herman.2013.Orga J. M. B. andP.M. deVries, Biermans, P.C.Klaasse T. Balke,G. Callaghan, patens under field conditions. Oecologia, 74:321-329. 74:321-329. conditions.Oecologia, patensunderfield fic coast of . fic coastof illo, B. Gallego-Tévar, E. Figuer illo, E. B. Gallego-Tévar, in the Iberian and S. maritimainthe Peninsula.Diversity ren, E. Martini, W. Vandenbruwaene, D. P. Vandenbruwaene, Martini,W. E. ren, A. Salmon, andM.Ainouc A. Salmon, ness. 2003. Inclusion 2003. offacilitationintoecological ness. oenergy. Journal of Journal oenergy. e study on three intertidal plant species. species. plant intertidal onthree e study rlogging responses in dune, swale and marsh swaleand responses indune, rlogging n, P. van Steeg, F. Dekker, J. vandeKoppel, J. F. P.vanSteeg, Dekker, n, high ecological amplitude between exotic exotic between high amplitude ecological nism traitsdeterminethestrength scale- of nism Ecology and Evolution 8:4992– and Evolution Ecology oa, B. Grewell, D.Vallet,H. Grewell, B. oa, Experimental Botany Experimental Spartina densiflora he. 2018. Lowhe. 2018. genetic Hacker, S. D., and M. N. Dethier. 2006.Comm S.D.,andM.N. Hacker, evolution in Acentury andA.F. of Raybould. D.F. 1991. Marshall, Gray, A.J., Chen, Gan, X.,Y.Cai,C.Choi,Z. Ma,J. RousseauH B.J., B., Grewell, Gallego- Tévar, D.K.Lee,a Friesen, P.C.,M.Peixoto, Ayres, A. D. Fortune, P.M.,K.Schierenbeck, Elliott, S.1821.Asketch ofSouth ofthebotany Hacker. 2005.Physical Dethier, M.N.andS.D. Davy.saltmarshLatin America.Pages179– Costa, C.B., 1992.Coastal communitiesof andA.J. .

Accepted .Oikos 113:279–286. marine impactsfor differing 21:1–62. Research anglica. AdvancesinEcological Estuarine, international importance. alterniflora inPlantEcology,. Perspectives hybrids maritimaandinvasive between Spartina native SpartinadensiflorainSouthwest 2019.Geneticstructureof Aïnouche,M.,Castillo,Salmon, A.,Figueroa, J.M. E., Journa climates. for cooltemperate Spartina Article 17:4304–4316. MolecularEcology context. invasive The enigmatic USA. marsh gra invasion ofanestuarine York, USA. 212 in U.Seeliger, editor. Coastal plant communitiesLatin of America. Academic Press, New pectinata onspring wetlandof birdcommunities atChongming aChinese Dongtan, (prairie cordgrass) and cordgrass) (prairie Spartina densiflora: ahistorySpartina inapolyploidy hybridizations of ss. Ecological Applications 15:1273–1283. ss. Ecological l of Experimental Botany 66: 4403–4413. Botany 66:4403–4413. l ofExperimental Coastal and Shelf Science 83:211–218. Science83:211–218. CoastalandShelf Evolution and Systematics 37:26-38. Systematics Evolutionand 37:26-38. and B. Li. 2009. Potentia nd R. F. Sage. 2015. Sub-zero cold tolerance of toleranceof nd R.F.Sage.cold 2015.Sub-zero Miscanthus ., Keller, J. Ainouche,A ., Keller,J. Bortolus, O. Clatrice, and M. L. 2008. O.Clatrice,andM. Ainouche. Bortolus, unity modification by a grass invaderhas grass unity a modificationby Carolina and Georgia. Schenck J.R., Charleston, J.R., Schenck Carolina andGeorgia. factors vs.bioticresistanceincontrollingfactors the × giganteus l impacts ofinvasive : candidate bioenergy crops bioenergy crops : candidate ., Lima, S., O.,Dréano, ., Spartina Spartina Spartina

Luque, G. M., C. Bellard, C.Bertelsmeier,Luque,G. M., C.Bellard, E. B. H.Wu, Zhao, J. Gan, Z.Y. Chen,X.J. L. H. Zhang, Chen,Q.Wang, Liao,B.,Li, X.D. C.H. E.D.Grosholz. and Levin,L.A., C.Neira, D.M.Richardson.201 and C.,P.Pyšek,Kueffer, B., Infante-Izquierdo, M.D., Gallego-Tévar, Sanchez Georgi Interactions between B. 1979. Haines, E. Dethier. 2009.Diff S.D.,andM.N. Hacker, .

Accepted16:981–985. i oftheworld’sworst Courchamp. 2014.The100th Engineering 35:511–520. Ecological effects. andecosystem status current of anoverview China: estuary, River Yangtze K.Chen. 2009. andJ. F.I.L. Jiang, Ma,X. Cheng, Z. J. 87:419–432. function. Ecology 633. multi-site studies,focusedme https://doi.org/10.1007/s00606-019-01591-5 ). subfamily densiflora (Poaceae, Article fromnative hybrids differentiation interspecific ofnew Castillo, MunozJ.M., Rodriguez, 2019.Mor A.F. USA. NewYork, Press, Systems. Plenum Pagesparadigm. 35–46 Berlin, Germany. Perspectives,Springer-Verlag, Conservation andManagement Pages 375–385 torestorationpotential. context community impactand modifying of significance invaders: in G. Rilov and J. Crooks, editors. Marine Bioinvasions: Crooks,editors. Ecology, Marine andJ. G.Rilov in R. J. Livingston, R.J. Ecological editor. ta-analysis andinvasionsyndromes.ta-analysis 200:615– Phytologist New 2006. Invasive cordgrass modifies wetlandtrophic Invasive modifies 2006. cordgrass ering consequences ofremoving ecosystem– consequences ering Bonnaud, P. Genovesi, D. Simberloff, and F. P.Genovesi, D.Simberloff,and Bonnaud, a salt marshes and coastal waters: A changing waters: Achanging salt marshesandcoastal a 3. Integrative 3. modelsystems, invasionscience: Plant Systematics and Evolution. Systematics andEvolution. Plant phological and anatomic Gullon, E., Nieva, F. J. J., Grewell, B.J., Grewell,B.J., J., Gullon, E.,Nieva,F. J. nvasive alien species. Biological Invasions Biological nvasive alienspecies. Spartina maritimaandinvasive Spartina alterniflora Processes inCoastalandMarine Processes al evidencesupports invasionsinthe S. Mendelssohn, I. T.Morris.2000.Eco- Mendelssohn, A.,andJ. Mobberley, D. G. 1956. Taxonomy and distribution of the genusand distributionofthe Taxonomy Mobberley, D.G.1956. and W. I. McKee L. Mendelssohn, A.,K. H.Wiersema.2015.Guidelines Redhead,S.A.,and J. McNeill, J., Morris, J. T. 1984. Effects of oxygen and salinity on ammonium uptakeby andsalinity ofoxygen T.1984.Effects Morris, J. Herrera Arrieta, and J. M. andJ. Arrieta, Herrera Peterson, P.M.,K.Romaschenko,Y. Pennings, S.C.,andM.D.Bertne S.A.Redhead,and McNeil, J., T.,P.V.Sundareshwar, C.T.Nietch, Morris, J. . Spartina alterniflora Massachusetts, USA. Massachusetts, USA. in Concepts andControversies alterniflora 64:1-4. Taxon reject names. Journal ofScience30:471–574. Journal conserve Massachusetts, USA. Associates, Sunderland, Community Sinauer M.E.Hay, editors.Marine Bertness, S.D.Gainesand Ecology. 64:1–4. Taxon reject names. coastal wetlandstorising Ecology 78:87–98. Loisel and Accepted 63:1373–1374. Chloridoideae: Sporobolinae ). Taxon Article Sporobolus Spartina patens :metabolic adaptation Loisel. Pages 59–80 against Spartina sea level. Ecology 83:2869-2877. sea level.Ecology 83:2869-2877. (Aiton) Muhl. Journal of Expe (Aiton)Muhl.Journal J. H. Wiersema. 2015.Guidelines H.Wiersema. J. Tidal Marsh Ecology. Kluwer Kluwer TidalMarshEcology. ss. 2001.Saltmarshco , H. Patrick. 1981. Oxygen deficiency inSpartina deficiency 1981.Oxygen H. Patrick. to anoxia. Science 214:439-441. Science214:439-441. to anoxia. in M. P. Weinstein and D. A. Kreeger editors. andD.A.Kreeger Weinstein M.P. B. Kjerfve, and D. R. Cahoon. 2002.Responsesof and D.R.Cahoon. Kjerfve, B. physiological controls on the productivity of controlsonthe physiological , Ponceletia mmunities. Pages289–316in rimental and , and Saarela. 2014a. Proposalto Saarela.2014a. Academic Publishers, Boston, Publishers, Academic Spartina for proposals to conserve or toconserve or for proposals Heleochloa for proposals to conserve or toconserveor for proposals . Iowa State College College Iowa State . Spartina alterniflora ( Poaceae: Poaceae: M.

Rilov, G., and J. Crooks, editors. 2009.Marine Rilov, G.,andJ. Silander, J. A.1979.Microevolu Silander, J. vandeKoppel,G. Ruessink, Bouma, vanBelzen, Z. J. Zhu, T.J. J. C. C.,O.Gourgue, Schwarz, andnative synopsis M.2012.Taxonomic ofinvasive Saarela, J. Rousseau-Gueutin, M.,S.Bellot, G.E.Marti Gray,Raybould,and R.T. F., A.J. A. 2013.China´s planis“overkill”.Nature499:392–393. Qiu, J. cordgrass Pergl, J. Pyšek, V.Jaro P., D.M.Richardson, Amolecular M.Saarela.2014b. andJ. Arrieta, Peterson, P.M.,K.Romaschenko,Y.Herrera .

Accepted660. life-history controlled by plant N. Claude,andS.Temmer 10:25–82. of including thefirstreport Northwest Chloridoideae) inthePacific Evolution93:5–16.and Phylogenetics and moleculardating. Molecular ofthehexaploid genome Salmon,A.K.Ainouc Coudouel, D.Naquin,A. Spring Management Perspectives. Article production andtheroleof purpurea biasesin invasionecology and taxonomic Sporobolinae classification of and newsubgeneric phylogeny in Poole Harbour: pattern of infection, effects onseed effects ofinfection, onSpartinaanglicainPooleHarbour:pattern ). Taxon 63:1212–1243. ). Taxon (Poaceae, Chloridoideae): Comparative analyses analyses Comparative Chloridoideae): Spartina maritima(Poaceae, Fusarium heterosporum man. 2018. Self-organization of a biogeomorphic landscape landscape ofabiogeomorphic man. 2018.Self-organization Spartina tion and clone structure in tion andclonestructure traits. Nature Geoscience 11:672–677. 11:672–677. Geoscience traits. Nature er-Verlag, Berlin, Germany. Germany. Berlin, er-Verlag, Claviceps ofClaviceps Thelong-termepidemic Clarke. 1998. xtownsendii for BritishColumbia, Canada. PhytoKeys (British Columbia,Washington andOregon), . Trends in Ecology and Evolution 23:237–244. in Ecology and . Trends n, J. Boutte, H. Chelaifa, O. Lima, S.Michon- Boutte, O. H. Chelaifa, n, J. šık, Z. Sixtová, and E. Weber. 2008. Geographical 2008.Geographical andE.Weber. Z. Sixtová, šık, Bioinvasions: Ecology,and Conservation he, and M. Ainouche. 2015. Thechloroplast andM.Ainouche.2015. he, Sporobolus . New Phytologist 138:497–505. . NewPhytologist 138:497–505. Spartina patens ( Poaceae: Chloridoidea: Poaceae: Spartina (Poaceae, (Poaceae, . Science203:658– Silliman, B., andA.Bortol Silliman, B. R.,andM.D.Bertness. 2002.Atr Silliman, B. R.,M.D.Bertness, andE.D. Strong,Ayres. 2013.Ecolog D.R.,andR. Ayres. 2009.Spartina A. Strong, D.R.,and IslesBritish ofthe (4th Flora Stace, C.2019.New Thomson, S.A.,R.L. S.T.Ahyong, M.Alon Pyle, va Bouma, J. Temmerman, S.,T.J. betweenecosystem Interactions andA.Bortolus.Sueiro M.C.,E.Schwindt,M.M.Mendez, 2013. of structure Sueiro M.C.,A.Bortolus, andE.Schwindt,2012.Theroleofthephysical . production. Proceedings of the National Academy of Sciences 99:10500–10505. 99:10500–10505. ofSciences of theNational Academy production. Proceedings marshes: marsh invertebrates eatmore marshes: marshinvertebrates University of California University of Impacts on editors.Human and E.D.Grosholz, thrive, and arrive, survive, UK. USA. Press,Berkeley, ofCalifornia University global perspective. marshes. A 16:e2005075. scienceisnecessary on 2018. Taxonomy based causes Herman. 2007.Vegetation i species Anative engineers: Annual ReviewofEcolog Accepteddensiflora Brong.A assemblages. instructuring macroinvertebrate Article us. 2003.UnderestimationofSpartina Press, California, USA. Press, California,USA. y 44:389–410. andSystematics sometimes hybridize. Pages3–22 ndirectly facilitates a non-native one. Acta Oecologica 51:11–16. Oecologicanon-native one.Acta 51:11–16. facilitatesa ndirectly n de Koppel, D. D. van der Wal, D.vanderWal, n deKoppel,D. channel erosion in a tidal landscape. Geology 35:631–634. inatidallandscape. Geology erosion channel than just detritus. Oikos101:549–554. thanjustdetritus. Grosholz, editors.2009.Humanimpactsonsalt Grosholz, ical and evolutionary misadventures of andevolutionary misadventures ical introductions and consequences insaltmarshes: introductionsandconsequences ophic cascaderegulate ophic for global conservation. PLoS Biology PLoS global for conservation. Ed). Cambridge University Press, Cambridge, Press,Cambridge, Ed). CambridgeUniversity Salt Marshes: AGlobal Perspective. Berkeley: so-Zarazaga, J. Ammirati, J. F. Araya, et al. F. Ammirati, Araya, J. so-Zarazaga, J. in productioninwestern Atlantic B. R.Silliman, M.D.Bertness, M. B. De Vries, and P. M. J. and P.M.J. De Vries, B. M. quatic Ecology 46:25–36. quatic Ecology 46:25–36. s saltmarshprimary Spartina Spartina . Yin, S., S. An, Q. Deng, J. Zhang, H. Ji, andX.Cheng. Zhang, 2015. H. Ji, J. Yin, S.,S.An,Q.Deng, L.Levin. C. R.,and Whitcraft Light-medi A. 2007. Liu,Wan, S.,P.Qin, J. andH.Zhou. 2009.The 1979Thenitrogen M.Teal. I.,Valiela, andJ. processe ecological I. Valiela, 2015.Marine F. H.,Barrie, R.,Gr J. Wiersema, Turland, N.J., Weinstein, M. P., and D. A. Kreeger, editors. M.P.,andD.A.Kreeger, Weinstein, . 81:192–199. salt marshplants.Ecology88:904–917. Ecology. Netherlands. Netherlands, Springer 656. Books. DOI https://doi.org/10.12705/Code.2018 July 2017. RegnumCongress Ve China, Shenzhen, Code)adopt (Shenzhen andplants algae, fungi, & Price,M.J. Smith, G.F.Prado, J., (eds.) Li, Marhold,K D.-Z., Knapp, S.,Kusber,W.-H., Acceptedimpact CH alterniflora Article 4 in China. Ecological Engineering 35:444–452. inChina. Ecological 35:444–452. Engineering andN 2 O fluxes from a salt marsh in eastern China. Ecological Engineering fromasaltmarshineasternChina.Ecological O fluxes s. Springer-Verlag, New York, USA. York, New s. Springer-Verlag, 2000. Concepts and Controversies inTidalMarsh 2000. ConceptsandControversies budget of a salt marsh ecosystem. Nature280:652– ecosystem. ofasaltmarsh budget 2018. International 2018. Code positive and negative effects of exotic andnegativeeffectsofexotic positive euter, W., Hawksworth, D. L., P.S., Hawksworth,D. Herendeen, W., euter, ed by the Nineteenth International Botanical International by theNineteenth ed ated regulation of the sediment ecosystem by ecosystem ated regulation ofthesediment ., May, ., Monro, A.M., J., T.W.,McNeill, getabile 159. Glashütten: Koeltz Botanical Koeltz getabile 159.Glashütten: Spartina alterniflora of Nomenclature for of Nomenclature invasions Spartina