Nomenclatural revision and syntaxonomical proposal for wetland peat vegetation in the Valdivian-Magellanian region
Abstract
Although numerous descriptive studies have been published throughout the 20th century on peat communities in the Southern Cone of South America, the nomenclature applied to the different units recognised often fails to take into account the proposals of previous authors. Our aim is to clarify the nomenclature by applying the rules of the ICPN to propose a current syntaxonomy of the class Myrteolo nummulariae-Sphagnetea magellanici. After a bibliographic compilation of possible peatland associations from the part of South America between parallels 40º and 56º south, under Temperate and Boreal macroclimates, we have collected around 30 names of phytosociological associations and alliances containing an abundance of homonyms and nomina nuda. Following ICPN standards, we have eliminated duplications and established priority names over others, lectotypified some names and rejected others as nomen ambiguum. We have arranged the floristic groups in the recognised associations in synthetic tables, with some nomina nudawhose characterisation we interpret as doubtful, as a proposed syntaxonomical organisation of the phytosociological class.The various communities recognised by the different authors who have worked in the territory can be divided into four physiognomic groups: 1. cushion bogs, 2. Sphagnumbogs 3. “montane tundra” bogs and 4. sedge-grass bogs. The syntaxonomic grouping we propose for the 13 associations recognised as nomenclaturally valid can be summarised as follows: a single class, Myrteolo nummulariae-Sphagnetea magellanici, is accepted, with a single order, Myrteolo-Sphagnetalia, and three alliances: Astelio pumilae-Oreobolion obtusanguli with seven associations (coinciding with type 1), Gaultherio-Sphagnion magellanici with four associations (type 2), and Abrotanello linearifoliae-Bolacion caespitosae with two associations (type 3). We contest the validity of around 22 names of syntaxa of different ranks, while acknowledging that more studies are required, particularly to define and delimit type 4 peatland communities
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165Lazaroa 38(2) 2017: 165-187 Lazaroa ISSN-e 1988-3307 http://dx.doi.org/10.5209/LAZA.56343 Nomenclatural revision and syntaxonomical proposal for wetland peat vegetationintheValdivian-Magellanianregion Javier Amigo1,CristinaSanMartín2,CarlosRamírez3&MiguelÁlvarez4 Received:24May2017/Accepted:2October2017 Abstract. Although numerous descriptive studies have been published throughout the 20th century on peat communities intheSouthernConeofSouthAmerica,thenomenclatureappliedtothedifferentunitsrecognisedoftenfailsto takeintoaccounttheproposalsofpreviousauthors.Ouraimistoclarifythenomenclaturebyapplyingtherules oftheICPNtoproposeacurrentsyntaxonomyoftheclassMyrteolo nummulariae-Sphagnetea magellanici. After a bibliographic compilation of possible peatland associations from the part of South America between parallels 40º and56ºsouth,underTemperateandBorealmacroclimates,wehavecollectedaround30namesofphytosociological associations and alliances containing an abundance of homonyms and nomina nuda.FollowingICPNstandards,we haveeliminatedduplicationsandestablishedprioritynamesoverothers,lectotypifiedsomenamesandrejectedothers as nomen ambiguum.Wehavearrangedthefloristicgroupsintherecognisedassociationsinsynthetictables,with some nomina nudawhosecharacterisationweinterpretasdoubtful,asaproposedsyntaxonomicalorganisationofthe phytosociological class. The various communities recognised by the different authors who have worked in the territory can be divided into fourphysiognomicgroups:1.cushionbogs,2.Sphagnumbogs3.“montanetundra”bogsand4.sedge-grassbogs.The syntaxonomic grouping we propose for the 13 associations recognised as nomenclaturally valid can be summarised asfollows:asingleclass,Myrteolo nummulariae-Sphagnetea magellanici,isaccepted,withasingleorder,MyrteoloSphagnetalia,andthreealliances:Astelio pumilae-Oreobolion obtusanguli with seven associations (coinciding with type1),Gaultherio-Sphagnion magellaniciwithfourassociations(type2),andAbrotanello linearifoliae-Bolacion caespitosaewithtwoassociations(type3).Wecontestthevalidityofaround22namesofsyntaxaofdifferentranks, whileacknowledgingthatmorestudiesarerequired,particularlytodefineanddelimittype4peatlandcommunities. Keywords: Argentina;Chile;cushionbogs;Sphagnumbogs;phytosociology;Myrteolo-Sphagnetea;peatlands. [es]Revisiónnomenclaturalypropuestasintaxonómicadelavegetacióndehumedales turbososdelaregiónValdiviano-Magallánica Resumen. AlolargodelsigloXXsehanrealizadonumerososestudiosdescriptivosdecomunidadesturbosaspor elCono-Sur sudamericano, pero confrecuenciala nomenclatura aplicadaalas distintas unidadesreconocidasno teníaencuentalaspropuestasdeautoresanteriores.NuestroobjetivohasidoclarificarlaNomenclaturaaplicandolas reglasdelCINFparaproponerunasintaxonomíadelaclaseMyrteolo nummulariae-Sphagnetea magellanici. Tras una recopilaciónbibliográficadeasociacionesdeturberasdelapartedeSudaméricacomprendidaentrelosparalelos40ºy 56ºsur,bajomacroclimasTempladoyBoreal,sehanreunidoalrededordeunos30nombresdeasociacionesyalianzas fitosociológicasentrelosqueabundanhomonimiasynomina nuda. AplicandolasnormasdelICPNsehaneliminado duplicidades,establecidonombresprioritariossobreotros,lectotipificadoalgunosnombresyrechazadoalgúnotro como nomen ambiguum.Losconjuntosflorísticosdelasasociacionesreconocidas,juntoconalgunasnomina nuda cuyacaracterizacióninterpretamoscomodudosa,sehanreunidoentablassintéticascomopropuestadeorganización sintaxonómicadelaclasefitosociológica. Ladiversidaddecomunidadesquehanreconocidolosdistintosautoresquehantrabajadoenesteterritoriosepuede repartiren4gruposfisionómicos:1.turberaspulvinadas,2.turberasesfagnosas3.turberasde“tundramontana”y 4.turberascipero-graminoides.Laagrupaciónsintaxonómicaqueproponemosparalas13asociacionesreconocidas comonomenclaturalmenteválidasseresumeen:unaúnicaclaseMyrteolo nummulariae-Sphagnetea magellanici,con unordenúnico,Myrteolo-Sphagnetalia,ytresalianzas:Astelio pumilae-Oreobolion obtusanguli con 7 asociaciones 1 LaboratoryofBotany,FacultyofFarmacy,UniversidadedeSantiagodeCompostela.E-15782SantiagodeCompostela(Spain). Email:javier[email protected] 2 EarthSciencesandHerbariumInstitute,FacultyofSciences,UniversidadAustraldeChile.Valdivia(Chile). 3 DepartmentofEcology,FacultyofBiologicalSciences,PontificiaUniversidadCatólicadeChile.Santiago(Chile). 4 INRES-Vegetationsökologie,Rheinische-FriedrichWilhelms-Universität,Bonn(Germany). ARTICLES
166 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 Introduction The southern part of South America delimited as the Valdivian-Magellanian region, according to the biogeographical proposal of Rivas-Martínez et al.(2011a),ischaracterized by its temperate (predominantly) or boreal (in the far south) macrobioclimates, as opposed to the precipitation deficit that is characteristic of the Mediterranean macrobioclimate in the adjoining Meso-Chilean-Patagonian region. This biogeographical region largely coincides with the area designated by classic authorsasthesub-Antarcticprovince(Cabrera & Willink, 1973), the sub-Antarctic subregion (Morrone, 2001) or Maritime Pacific South America (Josse et al.,2003).Someof best bioindicators that give the region its character are particularly the deciduous or perennial Nothofagusforests,andpeatlands.These peatlands have traditionally been the focus of interest due to their broad extension in TierradeFuego;alargenumberofstudieshave been done on the ecosystems in this southern part and in the Magellanian territory due to the logistical facilities in the towns of Ushuaia in Argentina,andPuntaArenasinChile.However,peatlandsofthesametypeandfloristiccontent extend to parallel 40º south in the northern partoftheValdivianbiogeographicalprovince due to the high levels of rainfall prevailing the lengthoftheChileanPacificcoast.Thisarea canbepermanentlyclassifiedwithinthehyperhumid or ultra-hyperhumid ombroclimatic belts,accordingtothebioclimaticclassification of Rivas-Martínez (1993, on line) and Rivas-Martínez et al.(2011b),andinterpreted fortheChileanterritoryinAmigo&Ramírez (1998)andLuebert&Pliscoff(2006). Although there are fewer studies on the peatlandsintheSouthernConethaninnorthern Europe, it is worth mentioning several worksthatproposedetailedclassificationsof peatland types. The most important of these were taken into account when studying the peatlandsinTierradeFuego,particularlythe work of Bonarelli (1917), and subsequentlytheclassicworksofRoivainen(1954)and Auer (1963) which examined in detail the variability in the floristic composition of these ecosystems.Theseclassificationswerebased mainly on structural and secondarily on floristicaspects,accordingtothebiotypesofthe dominant species. These classifications have continued to serve as the basis for more recent models that have set out to cover the whole range of peatlands in South American territory(Blanco&delaBalze,2004).Thereare also reference works from the 20th century with explanations of the ecology and the key parametersaffectingpeatlands;theseinclude theworksofMoore(1983)andPisano(1983), both of which lack vegetation data such as relevésorcompletefloristicchecklists.Vegetationdata,occasionallywithfloristicrelevés, can be found in studies from the same period onspecificareasintheChileanMagellanian territory: the contributions of Pisano (1971, 1972,1973,1977)andDollenz(1980,1982, 1986) are good examples. All this information basically focuses on MAG in bothArgentina and Chile. However,Oberdorfer(1960)wasthefirsttoformulate an integral syntaxonomic proposal: he collected his own relevés from the northern partof theVP (nearValdivia and the island ofChiloé,40º-42ºS),togetherwithsomelistedrelevésmadebyReiche(1907)insimilar areas. He highlighted the common elements between these relevés and others taken in the MAGterritory(54º-55ºS)basedonthework ofRoivainen(1954).Theseformedthebasis for the first formal proposals of phytosociological associations and alliances which were collectedinoneclass,Myrteolo-Sphagnetea, intendedtodescribethevegetationofChilean connombreválido(coincidentesconeltipo1),Gaultherio-Sphagnion magellanicicon4asociaciones(tipo2),y Abrotanello linearifoliae-Bolacion caespitosaecon2asociaciones(tipo3). Seargumentala invalidezdeunos22 nombresdesintaxonesdedistintorangoysereconocequehacenfaltamásestudiosenespecialparadefinirydelimitar las comunidades de turbera del tipo 4. Palabras clave: Argentina;Chile;turberapulvinada;turberadeSphagnum;fitosociología;Myrteolo-Sphagnetea;turbales. Nomenclature: Vascularplants:Zuloagaetal.(2009),withthesingleexceptionoftheconsiderationofthegenus Austroblechnum(Gasperet al., 2016).Bryophyta:ThePlantList(Anon.,2013).Lichens:Kirk&Cooper(2010). Abbreviations: ICPN: International Code of Phytosociological Nomenclature; MAG: combination of Temperate Magellanian and Antiboreal Magellanian biogeographic provinces; NVald: Northern Valdivian territory (40º-45º South);SVald:SouthernValdivianterritory(45º-47ºSouth);VP:Valdivianbiogeographicprovince.
167 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 temperate peatlands. New communities were published shortly after by Ramírez (1968) fromthesamenorthernVParea. Very soon afterwards Knapp (1966) published an outline of how to classify the peatlands in the “sub-Antarctic territory” which includedtheChileanregionofMagallanesand the Argentinian province of Santa Cruz; he proposedrecognizingasmanyasthreephytosociological classes: 1) Astelio-Oreoboletea, for hyper-humid peatlands with an absence of Sphagnum and which can be identified with whatwewillcallcushionbogs,dominatedby vascularplantspeciesgrowingintightclumps; 2) Rostkovio-Sphagnetea for peatlands with a dominance of Sphagnum spp. (particularly Sphagnum magellanicum) and characterized by a variety of bryophytic species; and 3) Carico-Calthetea, equating to wetlands with permanent waterlogging and incipient peat formation,whichcanbedescribedbytheautochthonous term mallines, containing sedge and grass vegetation. No association was formallypublishedin thiswork, butthe author cited some lists of plants he understood to be characteristic of the classes he proposed. The works dating from the last third of the 20th century failed to reach any unanimous criteria for interpreting and naming types of peatland;oneofthefewpointsofconsensus for practitioners of phytosociological methodology was to assume the name of the class Myrteolo-Sphagnetea to include the different communities. Probably the most intensive descriptiveworkwasonceagaindoneinMAG, where several associations and alliances were described in an international study conducted fromthePacifictotheAtlanticcoast(Roig et al., 1985); the proposals deriving from this work rejected various previously published associationnames,astheywereunderstoodto be communities that were not clearly delimitedorthat“shouldbeanalyzedwithmoreinformation”(Roig et al.,op.cit.: 482). In any case,allthenamesofnewsyntaxa that were formulated in this work lacked the formal rigourrequiredforthepublication,asbythenthe first International Code of Phytosociological Nomenclature (Barkman et al.,1976)wasalready in effect. Subsequent contributions differentiated Magellanianpeatlandsbasedontheirfloristic composition(Schwaar,1976,1981),ordemonstratedfloristicdifferentiationsrelatedtocontinentality gradients or the biogeochemical featuresofthepeat(Kleinebecker et al.,2007, 2008). New proposals have also been made in regard to peatland communities in the northern VP;thatis,informationsinthecoastalrange from Valdivia to Chiloé Island (San Martín et al., 1999, 2004; Ramírez et al., 2014a). Tosupplementthiswork,recentstudieshave been carried out on the peatlands in the region of Aysén (Teneb et al.,2008;Ramírez et al., 2014b,inpress;Rodríguez-Martínez,2015),a territoryofover100,000km2,hithertoverylittle studied and representing the biogeographicintermediarybetweenthenorthernVPand MAG.Thesecontributionshaveallservedas the basis for a list of peatland plant communitieswithnearly30associations,includinga proliferation of nomina nuda or homonyms. This highlights the need for a review of all the names described so far: the purpose is to serve as a guide to anyone wishing to apply phytosociological criteria to studies of peatlands in thisterritory,andtoclarifywhichassociations and alliances have been described previously and must be taken into account before new names are proposed. With all the validly publishedunitswewillultimatelyproposeafinal hierarchised syntaxonomical ordination. This is the aim of this work. Study area In addition to delimiting the biogeographic unit (Valdivian-Magellanian region), an ecological specificationmustbemadeastothetypeofwetlands whose communities we intend to discuss. This can be based on the distinctions expressed by Roig&Roig(2004),whodifferentiatemallines, peatlandsandmires,accordingtowhetherthe areasinquestionarepermanentlyoralmostpermanentlywaterlogged,withherbaceousvegetation and abundant organic matter which in certain circumstances may form peat (mallín); ecosystems where peat of various thicknesses has been produced and accumulated (bog or peatland);orareaswherepeatisbeingproduced andaccumulated,withaprogressivethickening of the organic deposit (mire). The communities westudygrowonpeatlandsormires,butthis delimitationisnotclearlydefinedbythevarious authors;althoughthereisanassumedcorrelationbetweentheEnglishterm“peatland”and theSpanishterm“turbal”;andbetween“mire” and“turbera”,thisdistinctionisnotuniversally accepted(León, 2012). Thus the name mallín
168 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 may even be used to describe the habitat of a particular community. With regard to the classification made by some Argentinian authors basedonthevegetation(Roig&Roig,op. cit.), we here study the communities framed within thedenominationof“peatlands”, whereasthe other two units, “meadows or mallines” and “peatmeadows”,includeawidearrayofformations containing a predominance of grasses and sedges that move progressively further from the concept expressed under the name of the class Myrteolo-Sphagnetea. They will thus only be cited incidentally at the end of this work. Although topography, and thus hydrogeomorphology, plays averyimportantrole in facilitating the installation of many peatlands (Rodríguez-Martínez,2015),onaglobalscale it appears evident that the factor most favouring peat formations is the regional climate regime: these formations are more frequent in areaswithapermanentlyrainyclimate(Málvarez et al.,2004).Inviewofthelargerain shadow thrown by the Andean mountains in the Southern Cone, as they contain the perturbationsoriginatingintheSouthernPacific (seeFigure1),itisthereforeonlynaturalto findagreaterextensionofpeatlandsinChilean territory, particularly in ombrotrophic territories or promoted by rain water. Chilean peatlands have been calculated to cover about10,470km2(Joosten&Clarke,2002), althoughChileanauthorsconsiderthismaybe anoverestimate(Schlatter&Schlatter,2004). No detailed estimates are available for Argentinianterritory,althoughtherearequantitative data referring only to the peatlands (excluding peat meadows and mallines) present in TierradeFuego,whichisalongwiththeFalkland Islands,themainArgentinianterritorywhere these ecosystems are found: an area of 500 km2 estimated by Bonarelli (1917). Figure1. SouthAmericanSouthernCone.A:BiogeographicalregionsaccordingtoRivas-Martínez et al. (2011a) classification: 13: Brazilian-Paranensean; 14: Chocoan; 15: Tropical South Andean; 16: HyperdeserticTropicalPacific;17:Pampean;18:MiddleChilean-Patagonian;19:Valdivian-Magellanian. Peatlandsinvolvedinthisstudybelongtounits19.1,19.2and19.3.B)Maximumannualrainfall(from Málvarez et al.,2004).
169 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 Biogeographical territories corresponding to the Valdivian-Magellanian region considered in this study can be delimited as follows: The VP extends from parallel 37º-38º to 47º-48ºsouth,and90%ofitsareacorresponds to Chile vs. 10% toArgentina.The greatest presence of peatlands in this province is associatedtohyperoceanicareas,whichcontinue to be abundant throughout the islands and the westernmostpartofthefjordsthatindentthe coastthroughtoCapeRaper.However,dueto their inaccessibility we have no data on these areas. Such hyperoceanic areas with important data on peatland vegetation are the coastal mountainrangebetweenthecitiesofValdivia andOsorno,ChiloéIslandandLosChonosIslands (40º-45º South). Onlyrecently,studiesonpeatlandsinthe region of Aysén in environments hyperhumid-humidhavebeendone.IntheResultsand Discussion section, data from territories between40º-45ºarecalled‘NorthernValdivian (NVald)’,whereasthoseinthebandbetween 45º-47ºarecalledsouthernValdivian(SVald). InadditiontotheVPandthechorological provinceofJuanFernández,whichhereiscompletely discarded due to the total absence of peatlands,theclassificationofRivas-Martínez et al. (2011a) distributes the remainder of the biogeographical region into three provinces: Temperate Magellanian,Antiboreal Magellanian and the Falkland Islands. With the exceptionofthedatafromtheseislands,theothers are not easy to assign to one or another province,asTierradeFuegoisdistributedbetween thefirstandsecond.Wethereforeoftenreferto datafromtheseprovincesassimply“Magellanian”(abbreviatedtoMAG)incomparison withthevegetationfromtheVP. Material & Methods We have examined the current and past bibliographycontainingfloristicinformationon possible peatland and bog communities. The authors’ proposals include numerous cases of overlapping floristic compositions which have given rise to a proliferation of names to describe combinations of species that could reasonably be interpreted as the same association,expressedwithdifferentaspects.Beforedeterminingwhether‘X’floristiccomposition expresses the reality of the association betterthananother‘Z’combination,wedecided to select the combinations whose publication has been validated according to the criteria stipulated in the third edition of the InternationalCodeofPhytosociologicalNomenclature (Weber et al.,2000;Izco&Del Arco, 2003). TheapplicationoftheICPNrules,andof article29binparticular,havealsorequiredthe modificationofsome association names that were illegitimate. With the set of nomenclaturallyacceptedassociations,andbasedonthe similaritiesbetweentheirfloristiccontents,we propose a syntaxonomical ordination of the communities they represent. Results & Discussion We describe the nomenclatural valid syntaxa in decreasing order of ordination in a syntaxonomical checklist. Acceptable names of syntaxa Startingwiththeupperranks,thefullnameis accepted for a single phytosociological class: Myrteolo nummulariae-Sphagnetea magellanici Oberdorfer 1960.Although Oberdorfer (1960) only discussed “Myrteolo-Sphagnetea” in his syntaxonomical checklist, there appears to be nodoubtaftertheauthor’sexplanationsthatthe name used to describe Sphagnum should be S. magellanicum. The name corresponding to the genus Myrteolamaybedebatable,asOberdorfer (op.cit.)repeatedlycited“Myrteola damaeonchii Berg”.Asnoted a few years later by Ramírez (1968), this was what was known by Chilean authors as M. barneoudiiBerg,andbyothersas M. nummularia (Poir.) Berg var. barneoudii (Berg) Kausel.InadetailedmonographonChileanmyrtles,Landrum(1988)clarifiedthatnoteventhe varietal rank was worth to distinguish and collected all the previously published names under the single binomial of Myrteola nummularia (Poiret) Berg;thisisthecriterionwefollowhere. Othersubsequentproposalscanbeconsidered.Knapp(1966)interpretedtherecognition of two classes: one Astelio-Oreoboletea for the peatlandsinhyperhumidterritoryidentifiable ascushionbogs,andaRostkovio-Sphagnetea for acidophilus Sphagnum bogs. To accept this separation at the level of the highest syntaxonomical rank it is necessary to recognise the priority of the alliance Astelio-Oreobolion
170 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 Oberdorfer1960(thepriorityisnotrecognised byKnappop. cit.),beforeusingitasthenomenclatural type for Astelio-Oreoboletalia and Astelio-Oreoboletea. As will be explained below, there are enough transitional situationsbetweenthesetwomajorpeatlandtypes (cushion and Sphagnum), with floristic convergencebetweenbothtypes,toruleoutthe option of two different classes. In addition, if the proposed class Astelio-Oreoboletea is to beaccepted,theothernon-cushionbogscannot be integrated in a syntaxon that bears the name of Rostkovio-Sphagnetea, as this group of communities would have priority eligibility to the name Myrteolo-Sphagnetea, making Rostkovio-Sphagnetea a younger homonym and therefore invalid. The option of a class Rostkovietea raised by Roig et al. (1985) is inappropriate in our view: in addition to being invalid nomenclaturally (as per article 5 oftheICPN),weunderstandthisproposalto havelittlefloristicsupportasthethreeassociations attributed to it by its authors appear to be a mixture of elements from bryophytic peatlands with characteristic species from the class Littorelletea australis of temporary lakes (seeDeil et al.,2011). The nomenclatural type of the single class is the order Myrteolo nummulariae-Sphagnetalia magellanici Oberdorfer 1960, the sole order whenthenameoftheclasswasfirstpublished. As described below, there have been some proposals,alsoinvalid(Roig et al.,1985),to includeanotherdifferentorder;asoutlinedin theConclusionssection,ouropinionistorecognise a single order. Thefirstpointtoconsiderindistinguishing alliances within this order is the two most widely verified models of peatlands that have been recognised by all authors studying peatland ecosystemsintheSouthernConeandwhichthey have sought to reflect in their classifications: Sphagnum bogs and cushion bogs. This can be interpreted as underlying the two alliances proposed by Oberdorfer (op. cit.), who sought a parallel between his personal observations in theNValdterritoryandthestudiesofRoivainen (1954) in MAG. Another classic author who studiedMAGterritoryindepthcoinedtheterm “Magellaniantundra”as oneof thefive major “biotic provinces” that can be recognised in MAG (Pisano, 1977), distinguishing within this category a cushion tundra and a Sphagnum tundra. The recognition of these ecosystems studied in Argentinian territory is open to a similar interpretation, as Roig (1998) collected only two alliances for his syntaxonomical checklist for Patagonia: one Sphagnion magellanici for Sphagnum bog communities; and the other Donation fascicularisforcushionbogcommunities, although both syntaxonomical names were ultimatelyinvalid,aswewillexplainbelow. Without venturing any proposals for communities, Kleinebecker et al. (2007) also coincided in recognising this dual model (cushion/ Sphagnum)alongawest-easttransectinMAG, which they clearly related to a continentality gradient and a decline in precipitation. Although this simplification excludes other possible physiognomic types worthy of consideration suchasgrassybogs(Pisano,op. cit.) or what is knownas“montanetundra”inMAG(Roig et al.,1985),itissufficienttostartbyrecognising associations that have been described since the mid-20th century to the present day. 1. Cushion bogs This is a particular group of communities as they represent a physiognomic type that is endemicinthesouthernhemisphere,occurringin the Valdivian-Magellanian region and also in TasmaniaandNewZealand,withwhichthey share genera but differ in their vicariant species (Godley, 1960; Wardle, 1991; Wardle et al., 2001;Ramírez et al.,2014a).Theirparticularity is the absolute dominance of vascular plants whichbuilddensecushions,andaverylow(in terms of biomass) participation of briophytes. CushionbogsareverycommonintheMagellaniantundra(Schmithüsen,1953;Pisano,1977, 1983) located around the Strait of Magellan in the coastal islands exposed to strong winds and abundant precipitation. Along the sinuous southwesterncoastofChiletheyextendnorthwardstotheChonosArchipelago,at44ºS(Álvarez et al.,2010);andthesecommunitiesare found even further northwards at higher altitudes,suchasat600ma.s.l.onChiloéIslandat 42ºS(Ruthsatz&Villagrán,1991)and1000m a.s.l.intheCordilleraPeladaat40ºS(Ramírez, 1968;SanMartín et al.,1999).Itshouldbenoted that the northernmost cushion bog communities were used as the reference for the earliest phytosociological descriptions due to their accessibility,andthenamesderivedfromthose studieshavepriorityaccordingtotheICPN.
171 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 Figure 2. Typical Donatia-cushion-bog which supports the weight of the botanist (doing his relevé) without deforming it. In the box we can see Donatia fascicularis, Astelia pumila and Drosera uniflora in dense folder.ChiloéIsland(LosLagos,Chile.42ºS). Table 1. Synthetic table of Astelio-Oreobolion and Gaultherio-Sphagnion. Floristic comparative among the recognized peatland associations included in Group 1 and 2. In addition,greycentralcolumnsshowthefloristiccompositionofthreecommunitiesdescribedasrelated togroup2butofuncertainsyntaxonomicposition(seetext).Theso-called“Sphagnetum magellanici” is also a nomen dubium. AssociationCode 1.1.1 1.1.2 1.2 1.3 1.4 1.5 1.6 1.7 Sph DoS Mar 2.1 2.2 2.3 2.4 N. of relevés 3 3 5 4 24 5 7 10 1 6 4 4 25 7 8 RelevéN. 1234567891011 12 13 14 15 Differentialsofassociations Schoenus rhynchosporoides 11II............ Schizaea fistulosa 2 2 . 1 . . III . . . . . . . . Gaultheria caespitosa 1..1.......3... Pinguicula chilensis .1............. Gentianella magellanica . . V............ Chusquea montana . . III 4 IV ......1... Racomitrium lanuginosum . . V1 I . . . . II . . . . . Marsippospermum philippi ...4I.......... Baccharis patagonica ....III.....1...I Schoenus antarcticus ....V.V. 1 V..... Lepidothamnus fonkii . 1 . . . VII . . V1 . . . I Austrolycopodium confertum .....II...I..... Senecio smithii .....II......... Schoenus andinus .....IV ..1...... Abrotanella linearifolia ......III........ Gunnera lobata . . . . . . II III 1 III 2 . . . . Caltha dioneifolia ......IV III 1 . . . . . . Dicranoloma hariotii .......V. . . . I III . Drapetes muscosus . . . . . . . II . . . . I III . Chorisodontium aciphyllum .......IV . . . . II I . Polytrichum alpestre ............III.. Juncus scheuchzerioides ........+...+.. Carex camptoglochin . . . . . . . II 1 . . 1 . . II
172 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 Gaultherio-Sphagnion magellanici Sphagnum magellanicum 22......5IV 1 4 V V V Carex magellanica 3 . . . . . . I 2 I 2 2 II III III Nanodea muscosa . . . . . . . II 2 I 1 . III V IV Sphagnum fimbriatum . . . . . . . . 1 . 1 . II I . Rostkovia magellanica ........1...IIII. Carex canescens ............+. I Astelio-Oreobolion obtusanguli Oreobolus obtusangulus 3 3 V2 . . IV V 1V2 . . . . Astelia pumila . 3 V4IV .V IV 2V. 1 . I . Donatia fascicularis . 3 V.IV V V IV 2V..... Tapeinia pumila . 1 III . I . IV III 2 . . . . . . Gaimardia australis . 1 V1+.IV III 1 V..... Tribeles australis .1.1.. .IV ....... Myrteolo-Sphagnetea magellanici Myrteola nummularia 2 3 V4 I . IV V 2 II 4 4 . IV IV Drosera uniflora . 3 V. II IV V II 2 V. 3 . I . Gaultheria pumila . . . 3 II III I II 3 IV 1 . IV III IV Acaena pumila . 1 . 1 . . III IV 1V2 . . IV . Tetroncium magellanicum . 2 . 1 . . I III 2 V2 . I VII Marsippospermum grandiflorum . 1 . . . . IV II 2 I 4 . I II IV Carpha schoenoides . 3 . 1 . . . III 1 I 2 . . II . Gaultheria antarctica . . . . . II . V. . . 1 . III Caltha appendiculata ......IIV2V2 . . IV I Pinguicula antarctica . 1 . 1 . . . . . I . . . . . Perezia lactucoides . 1 . 1 . . . . 2 . 2 . . . II Perezia magellanica .......I+. . . . I I Wintero-Nothofagetea species Nothofagus antarctica 1 . IV 4+.VI 2 IV 1. III IV I Nothofagus betuloides . . . . II IV III I . V2 . . . II Austroblechnum penna-marina 2 . . 3 . . . . . I 3 1 . . IV Pilgerodendron uviferum . . . 1 + V ...V1 . . . . Berberis serrato-dentata ...2I.......... Drimys winteri ....+. . I +...... Lebetanthus myrsinites ......IV .......I Philesia magellanica . . . . . I I . . I 1 . . . . Companionspecies Empetrum rubrum 1 1 . . . . . II 3 I 2 . IV II IV Cladonia confusa . . IV 2........... Gaultheria mucronata . . . . . I . . 1 II 3 . . . II Cortaderia pilosa . . . . . . II . 1 . 1 . . . . Thamnolia vermicularis ......IV . . II . . . . . Gunnera magellanica . . . . . . II . 2 . 4 . . I I Cladonia vicaria . . . . . . . III . . . . II I . Chiliotrichum diffusum . . . . . . . I 1 . 1 . I III I Other species: Cladonia pycnoclada 2, Sticherus quadripartitus, Dicranoloma sp. and Campylopus sp. 1, in 1; Baccharis magellanica 4, Senecio acanthifolius and Sphagnum sp. 3, Fitzroya cupressoides 1, in 4; Nothofagus nitida and Olsynium junceum +in5;Lomatia ferrugineaIIIin6;Tepualia stipularisIIin7;Festuca thermarum, Cladonia laevigata, Chorisodontium magellanicum, Dicranoloma robustum and Sphagnum falcatulum Iin8;another17species:shrubby(2Berberis),grass-sedgetype (2 Carex, 1 Uncinia, 2 Festuca) and chamaephytics (2 Azorella,2Bolax)in9;Cladonia pycnoclada and Dicranoloma billardierei II,Racomitrium wiliiV,in10;.Juncus cyperoides1in12;Chorisodontium leucopterum and Juncus balticusI,Uncinia tenuis and Hymenophyllum peltatumII,Luzuriaga marginataIV, in 15. Relevésource:1: Schoeno rhynchosporoides-Oreoboletum obtusanguli.Oberdorfer1960,Tb.43B;2:‘Schoeno rhynchosporoidesOreoboletum obtusanguli’ReicheinOberdorfer1960,Tb.43B;3:Drosero uniflorae-Donatietum fascicularis.Ramírez1968,Tb. 1;4:Astelio pumilae-Marsippospermetum philippiRamírez1968,Tb.2;5:Chusqueo montanae-Schoenetum antarctici.Ramírez et al.2014b,Tb.6;6:Schoeno andini-Lepidothamnetum fonkiiRamírez et al.2014,Tb.VI;7:Donatio fascicularis-Schoenetum antarctici Roig et al.1985,Tb.48,gr.72(subDonatietum fascicularis);8:Caltho-Oreoboletum obtusanguliOberdorfer1960,Tb. 44,col.’e’;9:Sphagnetum magellanici nomen illegitimum et dubiumPisano1977,Tb.XXI;10:Donatio-Sphagnetum magellanici nomen nudum.Roig et al.1985:Tb.48,gr.74;11:Marsippospermetum grandiflori nomen nudum.Roig et al.1985,Tb.48,gr.76; 12: Gaultherio-Sphagnetum magellanici.Oberdorfer1960,Tb.43;13:Polytricho alpestris-Sphagnetum. Oberdorfer1960,Tb.44, col.’a’;14: Oberdorfer1960,Tb.44,col.’b’;15:Nanodeo muscosae-Sphagnetum magellanici.Roig et al.1985,Tb.48,gr.75(sub Carico-Sphagnetum).
173 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 1.1. Schoeno rhynchosporoides-Oreoboletum obtusanguliOberdorfer1960 Lectotypus hoc loco: Oberdorfer(1960),pag. 160,Tab.43,rel.282a. This is probably the oldest association proposed for NVald, and is supported on six clearly heterogeneous relevés, half of whichwerecollectedbyOberdorfer(1960), and the rest cited from Reiche (1907) in severalwesternlocalitiesinNVald,though mostarefromChiloéIsland.Table1shows both halves separately to highlight their differences, since they in fact indicate two distinct communities with only six species in common. The half of SchoenoOreoboletumobtainedfromOberdorfer(rel. 1inTable1)hasalownumberofspecies, whereastheotherhalfobtainedfromReiche (rel.2inTable1)hasmanymorespecies, especiallythemostrepresentativeinGroup 1. The main formal problem with the last threerelevés(Reiche’srelevés)isthatthey containonlylistsofspeciesoccurrence,with noinformationontheirabundance,soonly onerelevéfromthefirstthreecanbeused asatypeforthisassociation,evenaccepting thattheyreflectafloristicallypoorsituation. In spite of the different biotypes of the two species, the association name does not need to be inverted (as “Oreobolo-Schoenetum”), in a hypothetical application of Article 29b of the ICPN. Although the sedge Sch. rhynchosporoides is taller than Oreobolus obtusangulus and most of the species in the dense mat, the cover index assigned by Oberdorferinasinglerelevéwiththepresence of this sedge was 1.2. 1.2. Droserouniflorae-Donatietum fascicularis Ramírez1968nom. inv. propos. Holotypus:Ramírez(1968),pag.96,Tab.1,rel. 3 [sub Donatia fascicularis-Drosera uniflora Assoziation] Thisassociationwaspublishedwithfiverelevés collected in the Cordillera Pelada (40°S) in a typical hyperhumid cushion bog dominated by Donatia fascicularis with the addition of Drosera uniflora, Gaimardia australis and Tapeinia pumila. There is also a presence of Oreobolus obtusangulus and Astelia pumila, though with less abundance than in association 1.1 but with a higher number of species. Two species must be highlighted for their biogeographical significance in relation to NVald: Schoenus rhynchosporoides and Chusquea montana,thelatteratypicaltaxon of bogs usually cited as Chusquea nigricans. SanMartín et al. (1999) established the lectotypeforthisassociationandinvertedthename, since Donatia fascicularis always acts as the dominant species and produces the highest biomass,andD. uniflora is not located in a high stratum with respect to Donatia;theauthorsof this article hereby submit a proposal to formalise the nomenclatural inversion. A recent proposal concerned Donatio fascicularis-Oreoboletum obtusangulae (sic), for cushion bogs from the coastal range on ChiloéIslandat42°S(Ramírez et al.,2014b). Weconsiderithasnomajorfloristicdifferences with Drosero-Donatietum fascicularis but represents a facies with a dominance of Oreobolus obtusangulus,andisotherwisecompletelysimilarto1.2,includingthepresenceofthebryophyte Racomitrium lanuginosum. Therefore the name “Donatio-Oreoboletum” must be interpreted merely as a synonym. 1.3. Astelio pumilae-Marsippospermetum philippi Ramírez1968 Holotypus:Ramírez(1968),pag.98,Tab.2,rel.2 [sub Astelia pumila-Marsippospermum philippi Assoziation] ThisassociationwasdescribedbyRamírez (1968)withrelevéscollectedintheCordillera Pelada,andwasfurthertypifiedbySanMartín et al. (1999). The main differences with the previous communities are the dominance of Astelia pumila,andthegreaterimportanceof the sedge Marsippospermum philippi and the dwarf bamboo Chusquea montana. Both these species support the discrimination from other communities dominated by Astelia pumila but described in southern territories. The higher cover of Astelia pumila can be correlated with initial succession stages in the vegetation dynamic during the regeneration of Donatia fascicularisbogs,assuggestedbyRuthsatz& Villagrán (1991) on the basis of palynologicalstudies;itmayalsoberelatedtothesubstrate,whichisamixtureofpeatandsand(San Martín et al.,1999), possibly explaining the participation of some woody species such as
180 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 Table 2. Abrotanello-Bolacion. FloristiccompositionofthefourmainassociationspublishedbyRoig et al. (1985) as representatives ofpeatlandcommunitiesof‘Montanetundrabogs’.Associationsrepresentedincolumns1and2 wereacceptedandtypifiedinthispaper;associationsrepresentedincolumns3and4stillremain as nomen nudum(seethetext).AsitwasshowninTable1,thecoincidentfloraintegratingthe Sphagnetum magellanici Pisano 1977 is exposed in column Sph (rel. 5) to underline the mixture character of that association. AssocCode 3.1 3.2 3 4 Sph Nr. of relevés 9 8 5 6 1 RelevéN. 12345 Differentialsofassociations Cryptochila grandiflora IV . . . . Senecio trifurcatus V. . . 1 Azorella selago .IV . . 3 Austrolycopodium alboffii . III . . . Abrotanella trichoachaenia . I . . . Carex kingii . I . . . Conostomum pentastichum . . . II . Lepidothamnus fonkii ...V1 Abrotanello-Bolacion caespitosae Perezia magellanica VII IV V + Austrolycopodium confertum II II II V. Drapetes muscosus IIV I I . Bolax caespitosa IV V V . 2 Phyllacne uliginosa VII V. 1 Orthachne rariflora II IV . . . Azorella lycopodioides II I . . 2 Viola tridentata I I . . . Andreaea sp. .IV I . . Austrolycopodium magellanicum .I.I. Myrteolo-Sphagnetea species Oreobolus obtusangulus VII III IV 1 Gaultheria pumila V IV IV III 3 Gaimardia australis VIIV IV 1 Donatia fascicularis II I III III 2 Tapeinia pumila IV I III II 2 Caltha dioneifolia IV V IV III 1 Thamnolia vermicularis IV I II III . Carpha schoenoides I I II . 1 Schoenus antarcticus III . I IV 1 Pinguicula antarctica I . I I . Abrotanella linearifolia III II . . . Nanodea muscosa I . II . 2 Drosera uniflora II . . III 2 Rostkovia magellanica . II I . 1 Caltha appendiculata . II . III 2 Astelia pumila . . VII 2 Tribeles australis . II . . . Marsippospermum grandiflorum . . III . 2 Myrteola nummularia ...V2 Acaena pumila ...V1 Perezia lactucoides . . . II 2 Wintero-Nothofagetea species Nothofagus antarctica IV I . I 2 Embothrium coccineum . II I I . Nothofagus betuloides . I I V2 Philesia magellanica . I . III 1 Lomatia ferruginea ...V. Pilgerodendron uviferum . . . III 2 Berberis ilicifolia . . . I . Escallonia serrata . . . I . Drimys winteri . . . I + Lebetanthus myrsinites ...IV . Empetrum rubrum . . . I 3 Berberis microphylla . . . I 2
181 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 Companionspecies Cortaderia pilosa I . IV . 1 Gaultheria mucronata I . . I 1 Gunnera magellanica . . . II 2 Racomitrium lanuginosum . I . . . Baccharis patagonica . . II . . Juncus balticus . . . I . Relevésource:1:Senecio trifurcati-Bolacetum caespitosae.Roiget al.1985,Tb.52,gr.78 [sub Bolaco-Phyllacnetum uliginosiRoig,Dollenz&Méndez1985nom. nud.];2:Azorello selaginisBolacetum caespitosae.Roiget al.1985,Tb.52,gr.79 [sub Azorello-Phyllacnetum uliginosiRoig, Dollenz&Méndez1985nom. nud.];3:Astelio-Phyllacnetum uliginosiRoig,Dollenz&Méndez 1985 nom. nud. Roig& al.1985,Tb.52,gr.80;4:Lomatio-Dacrydietum fonkiiRoig,Dollenz& Méndez1985nom nud.Roiget al.1985,Tb.52,gr.81;5:“Sphagnetum magellanici”.Pisano1977, Tb. XXI. [Including another 17 species]. Roig (1998) recognised six types of peatlands for the whole of the Argentinian part of Patagonia, two of which he called “Marsippospermum grandiflorum” peatlands and “waterlogged peatlands with Schoenus andinus and Carpha schoenoides”. Although he did not describe any floristic composition orrelevés,hisintentionwastoexpresscertain repeated physiognomic models. In the collection byMartínez-Carretero(2004),gradualfloristic changes can be observed in Tierra de Fuego from carpets of Astelia pumila and Sphagnum bogs of Sphagnum magellanicum through to various moist peat meadows with the participation of different sedges depending on the case: Carex gayana, C. magellanica, C. banksii, C. canescens, C. camptoglochin, and others. However this author omitted to give names to any of their communities. There was at least one proposal for a phytosociologicalassociationinChileanterritory to describe a type of sedge-grass peatland. Pisano himself (op. cit.) proposed Schoenetum antarctici,andgaveadescriptionaccompanied by one relevé. However, his single published relevé suffered from the same problem as his “Sphagnetum magellanici”,asexplainedabove: one relevé with 47 species of which over two thirdshaveanabundanceindexof“2”orhigher. We are therefore of the opinion that it should also be taken as a nomen dubium, and is not suitable to serve as the nomenclatural type for any association. Very recently we have highlighted the existenceofsedge-grassbogsintheSVald territory with the collection of 40 relevés from the region ofAysén, which we have named “community of Cortaderia pilosa and Schoenus andinus” with mean species richnessof14speciesperrelevé(Ramírez et al.,inpress). The detailed study of these peaty grass formationsthroughouttheSValdandparticularly MAGmayallowusinfuturetoarriveataphytosociologicaldefinitionofthistypeofvegetation,whosespeciesareoftenimbricatedwith communitiesfromGroups1-3. Rejected names of the syntaxa Table 3 shows a list of names of associations or higher rank syntaxa that have been discarded for reasons of invalidity. Although their conceptofcommunitymaybeacceptable(1, 4,11and12havebeenrecoveredtotypifynew names),theirnameisnot,andmusttherefore berejected. Oneoftheworkswiththelargestcontribution of nomenclatural proposals describing associations that ecologically differentiate the variouspeatlandsinMAG,Roig et al.(1985), fails to comply in all its proposals with the requirement of choosing a nomenclatural type (article5oftheICPN),sothenameofassociations proposed (1 to 4 and 9 to 14 in Table 3) are invalid. The name of association 5 is a clear case of nomen superfluum: Schwaar (1976) studied a community in Tierra de Fuego which he called Pernettyo-Sphagnetum magellanici, supported by 14 relevés whose floristic content in its typical subassociation barely exceeded six species; however, this composition is identicaltotheonepreviouslystudiedbyRoivainen (1954)andwhichOberdorfer(1960)proposed to call Polytricho alpestris-Sphagnetum. The problems with the names of associations 7 and 8 have already been explained as nomen dubium: their names should not be used,andnorshouldPisano’srelevés(1977), to typify new names.
182 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 The case of association 15 represents a flawedattempttousedatafromaclassicwork. SanMartín et al. (1999) proposed the name Astelio-Oreoboletum obtusangulae to designateatypeofcushionbogrecognizedinthe NVald territory. However they typified it by choosing a relevé from an association publishedbyOberdorfer(1960)underadifferent name: Schoeno-Oreoboletum.Giventhatthisis avalidlypublishedname(seeassociation1.1), “Astelio-Oreoboletum”isanomen superfluum whichmustberejectedbyapplicationofarticle29coftheICPN. AparticularcaseistheworkofRoivainen (1954),wholeftanextensivesampleoffloristic combinations found throughout numerous peatlands releved on 73 different sites in Tierra de FuegoinChileandArgentina;healsosupplied an even greater number of names for the different combinations he found on each site. There are examples of species combinations with nocommunityname,whileothersincontrast are designated with binomials, trinomials or tetranomials(“speciesA+speciesB+species C+speciesDSoziationen”).Weunderstandall these denominations, abbreviated as “-Soz.”, tobeinvalid,bytheapplicationof article 3d (whichreferstoPrincipleII)oftheICPN. Some proposed alliance names for groups of peatland associations must also be considered as nomina nudaandrejectedaccordingto ICPNspecifications: • Rostkovio-Sphagnion magellanici, proposedbyKnapp(1966)anddefendedin somemorerecentworks(SanMartín et al.,1999),hasneverhadavalidlypublished association that could be used as a type to support it. • Donation fascicularis has been proposed as an alliance name on two occasions. HoweverthefirstproposalbyRoig et al. (1985)combiningassociationsinMAG hadnotypeassociation;andthesecond, bySanMartín et al.(1999),isobviously invalid as it uses an identical name to a previous published name. • The name Astelion pumilae, proposed bySanMartín et al.(1999),wastypified on the association Astelio-Oreoboletum obtusangulae which is invalid, as explainedabove.Asaresultthetypification of this alliance is illegitimate. • The proposal of Sphagnion magellanici by Roig et al. (1985) is a case of the applicationofArticle31 of the ICPN, as thisnamewasusedmuchearlier(Kästner& Flössner,1933)forpeatlandintheNorthern Hemisphere.Othersyntaxaasthealliance Bolaco-Phyllachnion and its order BolacoPhyllachnetalia suffers the same problem asallnamespublishedbyRoig et al. (op. cit.):theabsenceoftypification. Conclusions Regardlessofthepossibleexistenceofpeatland vegetation units described based on someoftheirattributes,butwhichhavenot received an orthodox name according to the rulesoftheICPN,bycombiningallthephytosociological communities that have indeed beenvalidlynamed(Groups1-3),wecanarrive at a consistent taxonomical ordination to further the interpretation of this type of vegetationintheValdivian-Magellanianregion: There are communities in the so-called Magellanian tundra from parallel 40º to 56º souththatareidentifiableformaintaininga reasonably constant floristic composition in spite of covering such a broad distribution area. Although there is a much greater extension and abundance of these peatlands in MAG, there is no floristic contingent present in MAG that does not extend as far as NValdterritory,withtheexceptionofsome characteristic species from the associations in Group3(the“montanetundra”),agroupthat we interpret as genuinely and exclusively belongingtoMAG. AnalyzingGroups1and2incombination (which will subsequently be included as phytosociological alliances), the floristic variations that can be detected among associations in the same group are not due to biogeographicalbuttoclimatic,topographic and dynamic factors, to factors relating to thewatertable,orbecausetherelevéshave been taken in points of contact between communities. Very few species can be used for a biogeographical discrimination (Chusquea montanaexclusivetoNVald,and reciprocally Drapetes muscosus exclusive to MAG, see Zuloaga et al., 2009). We therefore do not consider it justifiable to establish more alliances within these groups; proposals such as “Donation fascicularis” or “Astelion pumilae” have been put forward more for reasons of circumstantial abundance of some of the dense carpet-forming species than for thediscriminationofspeciesofflora.
183 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 Table3.Namesofassociations,alliancesororderrejectedfollowingICPN.Therightcolumnspecifiesthe ICPNarticlesinvolved. N. Syntaxon Authors&publicationdate ICPN 1Donatietum fascicularis Roig et al. 1985 Art. 5 2Schoeno-Nothofagetum antarcticae Roig et al. 1985 Art. 5 3Donatio-Sphagnetum magellanici Roig et al. 1985 Art. 5 4Carico-Sphagnetum magellanici Roig et al. 1985 Art. 5 5Pernettyo-Sphagnetum magellanici Schwaar 1976 Art. 29c 6Marsippospermetum grandiflorae Pisano 1977 Art.2b,Art.7 7Sphagnetum magellanici Pisano 1977 Art.31,Art.37 8Schoenetum antarcticae Pisano 1977 Art. 37 9Marsippospermetum grandiflorae Roig et al. 1985 Art.5,Art.22 10 Cryptochiletum grandiflori Roig et al. 1985 Art. 5 11 Bolaco-Phyllachnetum uliginosi Roig et al. 1985 Art. 5 12 Azorello-Phyllachnetum uliginosi Roig et al. 1985 Art.3g,Art.5 13 Astelio-Phyllachnetum uliginosi Roig et al. 1985 Art. 5 14 Lomatio-Dacrydietum fonckii Roig et al.1985 Art. 5 15 Astelio-Oreoboletum obtusangulae SanMartín& al. 1999 Art.22,Art.29 16 Rostkovio-Sphagnion magellanici Knapp1966 Art.2b,Art.8 17 Donation fascicularis Roig et al. 1985 Art. 5 18 Bolaco-Phyllachnion Roig et al. 1985 Art. 5 19 Donation fascicularis SanMartín et al. 1999 Art. 31 20 Astelion pumilae SanMartín et al. 1999 Art. 17 21 Sphagnion magellanici Roig et al.1985 Art. 31 22 Bolaco-Phyllachnetalia Roig et al. 1985 Art. 5 The variety of ecological processes that converge in the formation and evolution of peatlands frequently produces different morphological types deriving largely from the abundanceordominanceofaparticularspecies; this has caused some botanists to promote association names based on the prominence of this circumstantially dominant species. This is particularly true in the case of cushion bogs,whereassociationshave beenidentified such as Oreoboletum, Donatietum or Astelietum, depending on the abundance of any of these three species (the three genera Oreobolus, Donatia and Astelia are monospecific in this territory); however, Marsippospermetum grandiflori or Lepidothamnetum fonkii have also been proposed due to the abundance of these two species in various circumstances. The dominance of certain species is often simply a question of changes in the depth of thewatertable,thethicknessofthepeatlayer, the dynamic phase of evolution towards woody communities, and so on. In these cases it is advisable to make a closer study of the water and nutritional dynamic of these peatlands and interpret any notable proliferation of certain species as simple variants or facies of some already known association. Syntaxonomical checklist Wesummarizethesetofcommunitiesthatwe havesofarconfirmedasvalidlypublishedin a syntaxonomical ordination (Appendix 1). BasedonthefloristiccontentsoftheassociationsshowninTables1and2,itcanbeconcluded that: It is reasonable to establish three phytosociological alliances with each one of the three groups of the previously explained associations. We excluderecognitionofanycommunityinGroup4 while awaiting more data from the whole of the VPterritory. Ingroup3werecognizetwoclearlydifferentiatedassociations,althoughweincludeanother twoinTable2,asnomina nuda,whoseidentity requirescarefulstudy.Weconsidertheirdifferentiatingfloraappeartobesufficientforthisgroup, and therefore propose the alliance Abrotanello
184 Amigo, J.; San Martín, C.: Ramírez, C. & Álvarez, M. Lazaroa 38(2) 2017: 165-187 linearifoliae-Bolacion caespitosae all. nova hoc loco [Syn. Bolaco-Phyllachnion Roig et al. 1985 nom. nud. p.p.]. Typus nominis: Azorello selaginis-Bolacetum caespitosae Amigo et al. ass. nova.Characteristicsofthealliance:Perezia magellanica, Drapetes muscosus, Austrolycopodium confertum, Bolax caespitosa, Phyllachne uliginosa, Azorella lycopodioides and Orthachne rariflora. Group 2 contains the Sphagnum associations in which a massive coverage of Sphagnum tends to serve as the primary selective criterion;thishassometimesledtotheinterpretationof“Sphagnetum”forpeatformations inverydifferentdynamicstages,orimbricated with cushion bogs. Table 1 contains a pair of communities published as nomina nuda, which have a floristically dissonant composition. For the four associations recognised we maintain the oldest validly published alliance name Gaultherio-Sphagnion magellanici Oberd. 1960. Typus nominis: GaultherioSphagnetum magellanici.Characteristicspecies: Sphagnum magellanicum, Carex magellanica, Nanodea muscosa, Sphagnum fimbriatum, Rostkovia magellanica, Chorisodontium aciphyllum, Polytrichum alpestre, Carex canescens, Gaultheria antarctica. The association that serves as the type for thisalliance,anobligatoryselectionunderthe termsoftheICPN(Art.20),isfloristicallyvery poor;thisshouldnotbeinterpretedasapretext for proposing as many new associations as there may be cases of Sphagnum bogs with the simple participation of a species that is not present in this association. The goal of a rational syntaxonomy is to address all the floristic variations that can contribute some biogeographical significance, and the participatingvascularfloradoesnotoffermany possibilities. A greater knowledge on species composition of the moss layer (bryophytes and lichens) of these peatlands would potentially make a more valuable contribution in this regardasindicatedbytheresultsofVillagra et al. (2009), and particularly by the recent contributionofQuilhot et al. (2012). Group 1, in which most associations have been recognised, also has the clearest floristic contingent, and is also the group for which most alliance names have been proposed, with the lack of validity explained above. It must have the oldest validly published name, namely Astelio pumilae-Oreobolion obtusanguli Oberd. 1960. Lectotypus nominis in hoc loco: Caltho-Oreoboletum obtusanguli Oberd. 1960. Characteristicspecies: Oreobolus obtusangulus, Astelia pumila, Donatia fascicularis, Tapeinia pumila,Gaimardia australis, Tribeles australis, Caltha dioneifolia. The set of three alliances, given the level of imbrication of the shared flora in all their associations,maybeintegratedinthesingleorder Myrteolo-Sphagnetalia magellanici. Lectotypus nominis in hoc loco: Astelio pumilae-Oreobolion obtusanguli Oberd.1960.Characteristicspecies: Acaena pumila, Caltha appendiculata, Carpha schoenoides, Drosera uniflora, Gaultheria pumila, Marsippospermum grandiflorum, Myrteola nummularia, Perezia lactucoides, Pinguicula antarctica, Tetroncium magellanicum. Weconsidertherestilltobeinsufficientinformation to accept the proposal of a second order suchas“Bolaco-Phyllachnetalia nomen nudum” fortheformationsweincludeinGroup3.Among otherreasons,acleardistinctionmustbemade between these moderately peat-forming associations of Abrotanello-Bolacion caespitosae and the nearby associations in clearly cryorophilous communities in the class Empetro-Bolacetea (Roig et al.,1985). Acknowledgments We would like to thank Prof. Jesús Izco (SantiagodeCompostela)forhishelpwith some nomenclatural aspects and with the proper implementation of the ICPN; L.G. Orellanaforthepreparationofmaps,andP. Brooke-Turner for the linguistic revision of the manuscript. References Álvarez,M.,SanMartín,C.,Novoa,C.,Toledo,G.&Ramírez,C.2010.Diversidadflorística,vegetacional ydehábitatsenelarchipiélagodeLosChonos(RegióndeAisén,Chile).An.Inst.Patagonia 38: 34-55. Amigo,J.&Ramírez,C.1998.AbioclimaticclassificationofChile:woodlandcommunitiesinthetemperate zone.PlantEcol.136:9-26.
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