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Home , Spartina, Spartina densiflora, Spartina patens, Sporobolus, Sporobolus alterniflorus

This article isprotected Allrights bycopyright. reserved. 10.1002/ecy.02863 version andtheVersi this between differences through the copyediting, typesetting, pagination forpublication accepted This articlehasbeen 3 Sydney, Australia SouthWales, New 2 (U9120ACD), Chubut, Ecosistemas Continentales 1 S. Weis Sytsma Salmon Pennings Muñoz-Rodríguez Hughes Figueroa M. Castillo Bertness Mark D. Alejandro Bortolus Running Supporting Head: 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. 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 (IPEEC), CONICET, Blvd. Brown 2915, Puerto Madryn (IPEEC),Blvd. Brown2915,Puerto CONICET, Spartina 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 and undergone full peer review buthasnotbeen review undergone fullpeer and on of Record. Please cite of Record. Please on and proofreading process, which may lead to 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 this article this as doi: 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 , This article isprotected Allrights bycopyright. reserved. LA, USA 20 19 USA 18 17 16 15 14 13 12 11 10 Lisbon, Portugal Lisbon, CampoGrande, 9 8 Department ofPhysical Geosciences, Faculty of and TheNetherlands; 140,4400ACYerseke, P.O.Box University, Utrecht (NIOZ), 7 6 5 042 Campus deBeaulieu, 35 4 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 Centre for and Hydrology, Ed 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 ofEnvironm Ri Universidade Federal do Departamento de BiologíayUniversidad Vegetal Ecología, de Sevilla, Spain Sevilla, Maritime Studies Program, Williams College Article and Environmental Science, Northeas and EnvironmentalScience, Conservancy, CA, USA Conservancy, CA,USA ental Science Policy, and University 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 inburgh Research Station,Scotland, UK Research inburgh Geography, Utrecht University, The Netherlands Netherlands The Utrecht University, Geography, 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, tern University, Nahant, MA, USA MA,USA Nahant, University, tern of California, Davis, CA, USA Davis,CA,USA California, of iences of the University of of the University iences This article isprotected Allrights bycopyright. reserved. 33 32 SC, USA 31 USA NC, Beaufort, University, 30 Argentina 29 Matieland, SouthAfrica 28 USA Arcata, CA, 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 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 a 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 nd Conservation, Nicholas School nd Conservation, 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 Coastal andfor 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 of the Environment, Duke of theEnvironment, osystems, Collegeof This article isprotected Allrights bycopyright. reserved. 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 species have contributed substantially (and specieshavecontributed for over 200 years. We do not agree with the subjective arguments underlying the years. withthesubjectivearguments donotagree We forover200 asasubgenusthatcouldrivalthestriki proposed the creation of a large monophyletic genus monophyletic ofalarge proposedthecreation Spartina mail: [email protected] mail: [email protected] Sciences, Rutgers University, Newark, NJ, USA NJ, Newark, University, Sciences,Rutgers to cal invasions, environmental ma cal invasions,environmental . itive feedbackthatwillstrengthen ortium, Sandy Hook, New Jersey, USA USA Hook,NewJersey, ortium, Sandy ical Laboratory, Woods Hole, MA, USA Laboratory, Hole,MA, Woods ical We understand the importance ofboththeobjective understand We theimportance 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 Sporobolus taxonomic decisionswithan taxonomic , and Sporobolus. , including (among Spartina

This article isprotected Allrights bycopyright. reserved. a acceptedby was Their proposal genera. conserving proposed they nomenclatural priority, of retain the name priority ICBN The 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 inthisnomenclatureis principle al. 2018).The Inte the and detailedin namingThe are taxa rulesInternational for governed by the forPlant Association withabasicchromoso dodecaploid species ofthispolyploid the emergence understanding of the closest relative(sister) lineages of morphological, andchromosomenumb physiological that provides critical insights into (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 monophyletic ofalarge proposedthecreation in 1810. While Petersonetal.(2014 in1810.While a,b ). This comprehensive phylogenetic re Thiscomprehensivephylogenetic ). Spartina first published in 1789, that predates recognition anddescription firstpublishedin 1789, thatpredatesrecognition theevolutionaryhistory ofthe 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 newly combined monophyletic genusshould newly combinedmonophyletic ming the paraphyly of the grass subtribe ofthe theparaphyly ming Spartina , which isofcrucial importance for Sporobolus a ) recognized that the name name the that ) recognized er evolution. It also priority , (Calamovilfa, A. Gray), Hack.,True Calamovilfa against against search is an important contribution isanimportant search , with a formal base date of 1753 basedateof , withaformal Sporobolinae menclature (ICBN; Turland et Turland (ICBN; menclature Sporobolus Spartina , and provides a clear picture picture provides aclear Sporobolus aswellother , encompassing its its , encompassing , including (among Spartina Sporobolus had

This article isprotected Allrights bycopyright. reserved. Accepted andsociology. history,ecology, economics, environmen ecology, experimental biogeography, knowledge inmultipledisciplines,including 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 cl amonophyletic representing (Mobberley 1956), The genus multidisciplinary often atefforts, immense cost. legacy significant of will servetounderminethe Articleproposal tochange the rationale underlying (M changes” “disadvantageous nomenclatural Spartina and well-rooted scientific, history of cultural andrelevance socio-economic implicit inthe name MYA;Rousseau-Gueu sister cladesometimes12-20 (e.g. clades divergent highly encompasses genus that large extremely presents majordisadvant This proposal,however, name Sporobolus Th forPlantTaxonomy. International Association Spartina . The important criterion for reje . Theimportantcriterionfor Spartina would avoid taxonomic destabilization, wouldavoidtaxonomic notin would necessitate many new binomialco new wouldnecessitatemany Schreb. is a morphologically we isamorphologically Schreb. a ). Its species are abundant and widely and abundant speciesare Its ). Spartina 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 ction or conservation ofbotan ction or Spartina evolutionary biology, molecular biology, molecular biology, evolutionary acNeill etal.2015).Wedonotagree withthe acNeill lly (and itcontinuescontributing)lly toour advances achieved after decades of worldwide ofworldwide decades after achieved advances ages including not only the creation ofan creation ages including the notonly tal management, restoration ecology, invasion ecology, management, restoration tal e authors argued that conserving the name the thatconserving e authorsargued ade (Baumel et al. 2002, Fortune et al.2007, Fortune etal.2002, ade (Baumel ll-circumscribed grll-circumscribed tin et al. 2015), but also overlooking thelong tin etal.2015),butalso to 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 Spartina ical namesistoavoidof ical oup ofgrasses oup divergedfromits This article isprotected Allrights bycopyright. reserved. Accepted 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 figured andbotany inagriculture prominently inbiogeomorphology well asadvances 2005,Hacker andHacker biology (Dethier theroleof inourunderstanding of keyadvances Silliman andBertness Silliman 2002, Article ecology inecosystem debate up" processes envir of 2003), thefunctioning andKr 1981,Weinstein 1979, Mendelssohnetal. includingthrough thenursery trophicinteractions the global onoutwelling-pulsing paradigm eventsin integrative ecology, “connectivity” the biodegradation pollutants, oforganic the negative vs.positive interactions incommunityecology, debatesonthedy zonation, in inmodernecology, concepts andperspectives spec iconic This . and subtropical temperate The smoothcordgrass 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” in the colonial (Eliot 1821,Barlett (Eliot colonialAmericas in the 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

This article isprotected Allrights bycopyright. reserved. Accepted ofclonalpl (Bertness 1991),microevolution work foundational including in marineecology, cordgrass The saltmeadow al. 2015). Ayres(Li et 2009,Wan and et al.2009,Strong These i secondary productivity. ecosystems vast coastal-marine Asia (Qiu2013)andthe in recorded events ever alterniflora ,China, coast, SouthAmericanAtlantic aggressive non-native asan is widely regarded speciesisoneofth etal.2018).This Schwarz organism traitsinlandscapeformation Articleinterdisciplinary field of “BioGeoMorphology”, 2013). andduplicationinpl 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 anglica of Other species S. anglica , known as common cordgrass, is a model ofre , knownascommoncordgrass, isamodel and S. anglica Spartina S. maritima provided the evidence to reconstruct the toreconstruct theevidence provided (Raybould et al. 1998). It etal.1998). isalsocons (Raybould have also made critical contri havealsomade are responsible for some ofthe forsome areresponsible S. patens and the introduced American American introduced andthe and alteredcoastal geomorphology nvasions resulted in striking,nvasions resulted has long been central toa haslong been (Bouma et al. 2005, 2013, Temmerman etal.2007, al.2005,2013,Temmerman et (Bouma 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 ants (Ainouche et al. 2004, 2012, Kueffer etal. al. 2004,2012,Kueffer ants (Ainoucheet species in , the North American Pacific AmericanPacific North inEurope,the species al. 2009, Saarela 2012, Luqu 2009, Saarela2012, al. designed to understand coastal plant zonation plantzonation coastal designed to understand 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 idered a key model speciesinthe a key model idered long-term epidemic oftheergot fungus long-term epidemic largest continental-scale bioinvasion continental-scale largest S. alterniflora cascading socio-economic impacts socio-economic cascading number of important experiments experiments ofimportant number , biodiversity, and primary and and primary , biodiversity, ntific knowledge. S. (Gray etal.1991, (Gray x townsendii e etal.,2014,Yin ofthe Spartina S. This article isprotected Allrights bycopyright. reserved. AcceptedSpartina participa “Spartina Watchers” scientists. The USAPacific instance, coast,commonly for is patrolled by volunteers called environmentalists.In itsintroduced range, itis the general public wholive alongIn coastlines the world. range itsnative around itisbeloved by Importantly, studying by achieved were to soilanoxia,drainage,and al. 2008,Castilloet2010,2018).Substa Fortune et atunpr spreading aggressively before decades show andSpain USA oceans, withworkinthe its hybridseluc alsopermitted environment new typeofcoastal Article Theaustralcordgrass Sueiro etal.2012,2013). Pacific andtheSouthwes nati for conditions andtheconsequences physical with (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 S. foliosa populations.This hasbeenrepeated Spartina and S.densiflora is one of a small handful of speciesth isoneofasmallhandful tern Atlantic respectively (Levin tern Atlanticrespectively Spartina and Sage, 2015, Friesen etal. andSage,2015,Friesen ting inearly-detection activ idation of the complex 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 ing that may lay fallow for may fallowfor lay that introducedspecies ing S. densiflora ypes inbioenergy biomassproduction forwoody ve and introduced invertebrates along the North theNorth along ve andintroducedinvertebrates 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 non-nativespeciesfollowacross 2015). Manipulative experiments experiments 2015). Manipulative at is known by itssc at isknownby is described as akey dominantina isdescribedas s etal.2009).Thisspeciesand S. pectinata lus 2006,Ayres etal.2008, is currently used iscurrently used ientific nameto Spartina

This article isprotected Allrights bycopyright. reserved. Accepted 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 in Giventhenum asaTopic. “Sporobolus” Topic “Spa underthe 2019) showed4,626papers Spartina alterniflora using listingsshows no(zero) Articlealterniflora " 7,290usesof records 16,2019) accessedMay (https://scholar.google.com/, scientific intheprimary peer-reviewed decade changing of broadconsensus, adopted (https://www.tropicos.org/) unilaterally have Ge databasessuchas While theimpactson global distribution, increasing todiscussthe countries since 1990indifferent professi attended by Conference”, Sporobolus 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 onal scientists and environmenta onal scientistsand to Spartina Spartina ber of species in each genus, 17in ineach ber ofspecies Sporobolus alterniflorus Sporobolus that include taxonomic updates on includetaxonomic that Sporobolus Spartina human society, and possible solutions. andpossiblesolutions. human society, literature. On the cont Onthe literature. problems associatedwiththisplantgroup, its vs184for than rtina”, while 690 papers were found using were rtina”, while690papers was under review, two papers (Gallego- Tévar Tévar (Gallego- review, twopapers was under has not found support for nearly half a hasnotfoundsupportfornearly half Sporobolus since the genus was first described in1789. sincethegenuswas 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 This article isprotected Allrights bycopyright. reserved. Accepted with otherexperts. Peterson etal.(2014 Within anthropology. literature and a striking ofst biology range asacross aswell name name committee rulesonthisquestion.Thereisclearl genus asa orsu isused to whetheraname mol relationships derivedfrom not genus does subgenus as a employed or ArticleSporobolinae new subgeneric classification..." (2014 Spartina nomen while potentially further any restraining 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, Spartina b Spartina ) proposed retaining retaining ) proposed is a deeply-nested, strongly s isadeeply-nested, and subgeneric classification of classification andsubgeneric for over 200 years.name The forover200 thatcouldrivalthestri asasubgenus b ) supply good basis for (re)naming other (re)naming good basisfor ) supply likely to induce the acceptance of replacing replacing of the acceptance toinduce likely Spartina ecular-based (ormorphological-bas ecular-based (title of paper) and "A pr (titleofpaper)and this context, the thiscontext, phylogene published in 2019 retains thename publishedin2019 as a subgenus of asasubgenus upported clade (Peterson etal.2014 (Peterson clade upported 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 Spartina akeholders anddisciplines, including history, akeholders Sporobolus clatural discussions clatural y tosubsume the no rationalesocompelling king, global iconic history and use of the and useofthe globalking, iconichistory rings as a signature name in ecology and name inecology rings asasignature Sporobolus " (Table 2 legend)). Whether a name is aname Whether " (Table2legend)). oposed classification ofthe classification oposed Sporobolinae tic relationships providedbytic relationships ("A molecular phylogeny and phylogeny ("Amolecular ed) phylogenies. Decisionsas phylogenies. ed) and sound open debate. and soundopendebate. Spartina Spartina a,b clades in agreement agreement in clades for ). Petersonetal. Sporobolus after considering , This article isprotected Allrights bycopyright. reserved. Acceptedsuppliedthe excellent Editor-in-Chief who which they are affiliated.We are alsograteful deeply totwo anonymous reviewerswell as to theorgani reflect personal, anddonotnecessarily improving ourreasoningand encouraging ustoinitiate thisdebate. positions are The authors’ forsh UK andUruguay Spain, Argentina, USA, stude colleagues, many the to grateful are We Acknowledgments et al.2014 assubgenus designated clade well-knit distinct the that consider therefore, should continuetolinkscientists, citizensa impacts.Thename global 2019)withprofound (IPBES change innature marshes globally as well as contri Spartina Article impactinsomany suchanimmense ever provided d strengthen taxonomic positive feedbackthatwill scientific name such thatmayaccompany nomenclature formalized understandtheim indeed, wefully ge as a should notbeinterpreted but,ra nomenclatural viewpoints criti deserves question, whichwefeel insights. said,onthisrepor phylogenetic This thevalueofmolecula recognize unequivocallyWe , intimately tothe linked initial recognition of a , should simply and efficiently be treated as the genus asthe treated be efficiently , shouldsimply and s are hypotheses, ourhopeisthatby s arehypotheses, 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. insight that truly improved our work. insightwork. that truly improvedour nd environmentalmanagers nts and eight anonymous expert taxonomists from taxonomists eightnts and expert anonymous t we have focused on a on t wehavefocused 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 opening thisdebate opening 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 theglobe. across We, we will encourage we Spartina ntific fieldsashas ntific . Spartina haslinkedand by Peterson This article isprotected Allrights bycopyright. reserved. 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, Ferreira DeCarv J. Ainouche, M.,H.Chelaifa, 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 Phylogeny of hybridizing species from the genus speciesfromthe Phylogeny of hybridizing BritishGuiana1:5–10. of Agriculture of 713–719 native and D.R.Strong.Hybr 2008. USA. New York, 225–244 2012. Polyploid in evolution Spartina Ricou, andM.-T.Misset.2009.Hybridizati 484. Bi genomes. allopolyploid changes thataffect genomics: Cambridge. AcceptedEvolution 22:303–3014. Phylogenetics and Article S. foliosa () Biological Invasions 11:1159–1173. Biological (Poaceae) in Spartina anglica P.S.Soltis and Springer, D.E.Soltis,editors.Polyploidy Evolution. and Genome in San Francisco Bay, California, US Bay, inSanFrancisco California, Schreb. a natural model system for analyzing early evolutionary early analyzing system for model Schreb.anatural 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 Schreb. (Poaceae). Molecular Molecular (Poaceae). Schreb. (Poaceae) and (Poaceae) This article isprotected Allrights bycopyright. reserved. Low,Vries, E. G.Peralta M.B.Bouma, De T.J., T.Carlton, andE.Schwi Bortolus, A.,J. of 2009.Acharacterization L.Idaszkin. Bouza, andY. P. Bortolus,J. 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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 densiflora ofhybrids with 2010.Theproduction Figueroa. 66:4093–4095. Articlepopulations of 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. deVries, Biermans, P.C.Klaasse T. Balke,G. Callaghan, Spartina andnative 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, S. maritima s and salt deserts of the world. Interscience Pub. Inc., Interscience Pub. New oftheworld. s andsaltdeserts ren, E. Martini, W. Vandenbruwaene, D. P. Vandenbruwaene, Martini,W. E. ren, A. Salmon, andM.Ainouc A. Salmon, ness. 2003. Inclusion 2003. offacilitationintoecological ness. in the Iberian and inthe Peninsula. Diversity 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

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pectinata onspring wetlandof birdcommunities atChongming aChinese Dongtan, (prairie cordgrass) and cordgrass) (prairie Spartina maritima Spartina Spartina densiflora Spartina 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 andinvasive Miscanthus ltmarsh communities of Latin America.Pages179– ltmarsh communitiesof : ahistory inapolyploidy hybridizations of ., Keller, J. Ainouche,A ., Keller,J. Bortolus, O. Clatrice, and M. L. 2008. Ainouche. O.Clatrice,andM. Bortolus, stillo, J.M. 2019.Geneticstructureof stillo, J.M. 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 × Spartina densiflora giganteus l impacts ofinvasive : candidate bioenergy crops bioenergy crops : candidate ., Lima, S., O.,Dréano, ., inSouthwest Spartina Spartina Spartina

This article isprotected Allrights bycopyright. reserved. C.Bertelsmeier,Luque,G. M., C.Bellard, E. B. Zhao, H.Wu, J. Gan, Z.Y. Chen,X.J. L. H. Chen,Q.Wang, Zhang,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, Engineering 35:511–520. 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 densiflora fromnative hybrids differentiation interspecific ofnew Castillo, MunozJ.M., Rodriguez, 2019.Mor A.F. USA. NewYork, Press, Systems. Plenum Pagesparadigm. 35–46 Conservation andManagement Pages 375–385 impactand modifying of significance invaders: Accepted16:981–985. i oftheworld’sworst Courchamp. 2014.The100th Article (Poaceae, subfamily ). subfamily (Poaceae, in G. Rilov and J. Crooks,editors andJ. G.Rilov in R. J. Livingston, R.J. Ecological editor. ta-analysis andinvasionsyndromes.ta-analysis 200:615– Phytologist New Perspectives, Springer-Verlag, Berlin, Germany. Berlin, Germany. Perspectives,Springer-Verlag, 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 community context torestorationpotential. context community . Marine Bioinvasions:. Ecology, Marine 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 maritima Spartina alterniflora Processes inCoastalandMarine Processes andinvasive al evidencesupports invasionsinthe S. This article isprotected Allrights bycopyright. reserved. 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. on ammoniumuptakeby andsalinity ofoxygen T.1984.Effects Morris, J. genusand distributionofthe Taxonomy Mobberley, D.G.1956. I. T.Morris.2000.Eco- Mendelssohn, A.,andJ. and W. I. McKee L. Mendelssohn, A.,K. Redhead,S.A.,and McNeill, J., conserve 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 ofScience30:471–574. Journal Massachusetts, USA. in Concepts andControversies Spartina alterniflora alterniflora 64:1-4. Taxon reject names. AcceptedChloridoideae: Sporobolinae Article Sporobolus Spartina patens :metabolic adaptation Loisel. Pages 59–80 against Massachusetts, USA. Massachusetts, USA. 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. Spartina ). Taxon 63:1373–1374. ). Taxon J. H.Wiersema.2015.Guidelines J. Tidal Marsh Ecology. Kluwer Kluwer TidalMarshEcology. ss. 2001.Saltmarshco Herrera Arrieta, and J. M. andJ. Arrieta, Herrera , H. Patrick. 1981. Oxygen deficiency in 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. Pages289–316 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: Spartina in M.

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Accepted81:192–199. impact CH salt marshplants.Ecology88:904–917. Netherlands. Ecology. Netherlands, Springer alterniflora Article656. 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., 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. ConceptsandControve 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: Spartina alterniflora of Nomenclature for of Nomenclature rsies in TidalMarsh rsies Koeltz Botanical Koeltz invasions Spartina