Cryptogamie,Bryologie,2010,31(1):75-94 ©2010Adac.Tous droits réservés

Philonotiscapillaris Lindb.and P.arnellii Husn.; one ,twonames

Timo KOPONEN* &PekkaISOVIITA

BotanicalMuseum(bryology),P.O.Box7, FI-00014 University of Helsinki,Finland

(Received 10September2009,accepted 30 September2009)

Abstract – Philonotiscapillaris, described byS.O.Lindbergin1867,hasbeen variously treated. Some authors considered ittobeasmall formof P.marchica,but Lindberghimself combined itas P.fontana var. capillaris on the basisof its spreading perigonialleaves,which areerectin P.marchica. P.T.Husnotwasconfused byLindberg’ssolution and described the same taxon as P.arnellii in 1890.Manynewrelated speciesweresoon described,and for awhile some authors accepted both P.capillaris and P.arnellii.H.N.Dixon provisionally synonymised P.arnellii with P.capillaris in 1896,while L.Loeske (1906)and M.G.Dismier (1908) did thisdefinitively.Nonetheless,while NorthAmericanflorasand checklists unanimouslyusethe correctname P.capillaris, the name P.arnellii isstill used in almost all recentEuropeanflorasand checklists.The lattername hasuntil nowbeen cited from Husnot’s“MuscologiaGallica”(Jun-Jul1890),but the valid publication byHusnotin Revue Bryologique(Apr-Mai1890)antedatesthis.Hereweclarify,based on thatcorrect protologueand Husnot’sletters toS.O.Lindbergand V.F.Brotherus,whyHusnot described P.arnellii.

V.F.Brotherus /History /T.Husnot/S.O.Lindberg//Philonotisarnellii / Philonotiscapillaris /Nomenclature/Synonymy

INTRODUCTION

Thiscircumpolar,temperateand borealmoss wasdescribed and named Philonotiscapillaris byLindberg(1867)from Danishand Swedishmaterial; however,the validity of thatfirst publication hasbeen questioned. Crum etal. (1973)attributed the name toLindb. ex C.J.[sic] Hartm. (1871),while Crum& Anderson (1981) and Allen (2002)explained that Philonotiscapillaris Lindb. (1867)isa nomen nudum and thatthe name wasonlyvalidated byHartman(i.e. C.Hartman,1871). Alatername, Philonotisarnellii, waspublished byHusnot(1890a, b)on the basisof specimensfrom Sweden. P.arnellii Husn. hasbeen synonymized with P . capillaris bynumerous contemporary European(e.g. Dixon &Jameson,1896 provisionally; Dismier,1908,1910; Brotherus,1909,1923,1924;Mönkemeyer, 1927)and recent,especiallyAmerican,authors (Anderson etal.,1990; Allen, 2002),doubtless on taxonomicallygood and nomenclaturallycorrectgrounds.In spiteofthe unanimous acceptanceof P.capillaris byAmericanbryologists,the

*Correspondenceand reprints:[email protected] 76 T.Koponen &P.Isoviita name P.arnellii isstill used bythe majority of recentEuropeanauthors.The presentarticle aimstoelucidatethe reasonsforthisconfusing practiceand to encourage auniformusage on bothcontinents.

PHILONOTIS CAPILLARIS LINDB.

Inthe protologueof Philonotiscapillaris, Lindberg(1867)did notcite specimensaccurately,stating onlythatthe plantgrows in Denmarkand Sweden. The description of P.capillaris issimilarlyvery short.Inthe same paragraph Lindbergfirst statesthatthe moss named byhim as P.parvula [ nomen nudum ]in 1859 must beregarded asavariety of P.muehlenbergii (Schwägr.) Brid. and then continuesasfollows: “Ob aber Ph. Mühlenbergii von Ph. marchica hinlänglichveschieden sei,wage ich nochnichtzuentscheiden;esscheintmirjedochso. Desgleichen ist mirnocheine andereArt derselben Gattung etwasunklar,welche aufnacktem Boden Schwedensund Dänemarkswächst; sie ist beinahe haarfein mitäusserst schmalen Trieben und sehrausgezeichnet,stehtaberuntereuropäischen Formen deroben genannten amnächsten. Von dieser,welche ichinmeinem Mss.von 1865als Ph. capillaris beschrieben habe,besitzeichleidernur ein einzigesund unvollständigesFruchtexemplarund sehrwenig männliche,dagegen sind die sterilen weiblichen minderselten.” Lindberg’s(1867)characterisation of Philonotiscapillaris, “sie ist beinahe haarfein mitäusserst schmalen Trieben und sehrausgezeichnet”,isa sufficientdescription,and the name wasaccepted in “IndexMuscorum” (Wijk et al.,1967,1969) asthe basionymof P.fontana (Hedw.) Brid. var. capillaris (Lindb.) Lindb.Crosby etal. (1999) alsoattributed P.capillaris to“Lindberg,1867”,and itiscertainlynotabare nomen nudum .Nonetheless,the name isnotvalidly published if itisobvious that“itisnotaccepted bythe author[Lindberg]in the originalpublication” (see Art.34.1(a )ofthe Code,McNeill etal. 2006). Astothis nomenclaturalpointof view,insteadofdirectlypresenting hiscurrentopinion Lindbergstated thatin amanuscriptof 1865hehaddescribed the speciesas P.capillaris .Similarexpressionsof namesnewlyestablished in hisearlier manuscripts arenotunusualinLindberg’sprotologues.Hewasanabsolute defenderof the priority principle and approved nominanuda and otherinvalidly published namesaswell asmanuscriptnames(Isoviita, 1966:209). Lindberg’s referencetohis“Mss.von 1865” isintended toshowthatthe name P.capillaris gotpriority in 1865. Thereisno doubtthatLindbergstill accepted itin the 1867 publication. Onthe otherhand,hisslighttaxonomichesitation in 1867 concerning the somewhatunclear(“etwasunklar”) speciesstatus seemstobeduemerelyto him nothaving available aspecimen withcompletesporophytes, i.e. withold capsules.Inanyevent,the last sentenceofthe Article cited abovestates:“Art. 34.1(a )doesnotapplytonamespublished withaquestion markorother indication of taxonomicdoubt,yetaccepted bytheirauthor”. Thisprovision confirmsthatsuchahesitation alone cannotinvalidatethe 1867 publication. Even if Lindberg’s(1867)first publication isnotconsidered acceptable, Philonotiscapillaris wasvalidated as Bartramiacapillaris “S.O.Lindberg(in litt.)” byT.Jensen (1868) and,independently,as Philonotiscapillaris (Lindb. ex T.Jensen) Milde byMilde (1869). Inspiteofthe latternotciting T.Jensen (1868), underArt.33.3of the Code Milde’sname canberegarded asavalidlypublished combination based on Bartramiacapillaris .Furthermore,Milde alsoascribed Philonotiscapillaris Lindb.and P.arnellii Husn.;one moss,twonames77

P.capillaris toLindbergand evidentlyknewLindberg’s(1867)original publication. InhislettertoS.O.Lindberg(in the archivesof FinnishNational Library,see Koponen &Isoviita, 2005),dated in Breslau“d. 16.Febr.68.”,Milde wrote:“ Hylocomiumsubpinnatum und Philonotiscapillaris haben wirjetzt auch in Schlesien.” The citation Philonotiscapillaris Lindb. ex C.J.Hartm. or P.capillaris Lindb. ex Hartm.,frequentlyused byAmericanauthors,refers toC.Hartman’s (1871) treatmentin the 10thedition of the Scandinavianflorafounded byhis father,C.J.Hartman(1790-1849). Thispublication of P.capillaris hasno nomenclaturalstanding sinceamerehomotypicisonym(asdefined in Art.6 Note2of the 2006 Code) isinvolved. AfterHartman’s(1871) flora, and beforethe description of Philonotis arnellii (see below),differentopinionsof the status of P.capillaris werepublished. Intheirlist of all of the mossesof the world,Jaeger&Sauerbeck(1875) cited Lindberg(1867)withaquestion mark,and mentioned alsoMilde (1869). Schimper(1876)thoughtthatspecimensfrom Germanyand Scotland are differentfrom the Danishplant,whichisagracile formof P.marchica (Hedw.) Brid. Limpricht(1876)and Molendo (1875) considered P.capillaris tobeavariety of P.marchica, and Geheeb(1878) did notaccept P.capillaris asaspecies. Zetterstedt(1876)maintained the specificstatus and presented P.parvula Lindb. [ nom. inval.]asits synonym. Gravet(1883)marked P.capillaris Lindb.asan uncertain and insufficientlyknowntaxon.

PHILONOTIS ARNELLII HUSN.

Instandardindicessuchas“Indexbryologicus”(Paris,1896,1905) and “Indexmuscorum” (Wijk etal.,1967),aswell asearlyand laterfloras(e.g., Limpricht,1893; Warnstorf,1905;Möller,1925;C.Jensen,1939;Nyholm,1960, 1998;Lawton,1971;Smith,1978,2004),the name Philonotisarnellii iscited from the 9thpart (“livraison”) of Husnot’s“ MuscologiaGallica”(1890b). However,in the same yearHusnotpublished anarticle on dioicous speciesof Philonotis (Husnot,1890a)in RevueBryologique 17(3):42-47,and P.arnellii isdescribed alsointhatarticle. Ifpublished earlierthan“MuscologiaGallica”,the article in RevueBryologique constitutesthe realprotologue. Podp{ ra(1954) cited alsothe article in the “ Revue ”and Möller(1925) mentioned itin the text. InTL-2’sentry “ 3154. MuscologiaGallica”,Stafleu&Cowan(1979) state: “Datesbased on notesin Nat.Nov.,Hedwigia, Bot.Gaz.,Bot.Zeit,and Rev. Bryol.”. The dateTL-2providesforLivraison 9and forp. I-VIII of the entire volume including its reissued Plates1-10isJun-Jul1890,whereasall of the other 13parts havemoreprecisedates.Among the sourcesmentioned above,only Bot. Gaz .15:275(Oct1890), Nat.Nov .12:360 (2nd half of Aug1890),and Rev.Bryol. 17(4):60-61(ca Jul-Aug1890)canservefordating the 9thlivraison;and apparently itisthe latterfrom whichthe approximatedateJun-Jul1890hasbeen drawn. Inthe context involved,areviewof Husnot’snewcontribution,the pagesI-VIII arealso mentioned and the following information isprovided:“Lessouscripteurs recevront en même tempsquecettelivraison unnouveautirage des10premièresplanches…” (Husnot,1890c). Afterall,itisquitelogicalthatthe first reviewof Husnot’sbook waspublished in the journalfounded and edited byHusnothimself, Revue Bryologique ,and assoon asthe completebook became available. 78T.Koponen &P.Isoviita Philonotiscapillaris Lindb.and P.arnellii Husn.;one moss,twonames79

Fig. 1. P.T.Husnot’slettertoS.O.Lindberg. Originalinthe Archivesof the NationalLibrary of Finland. All lettercopiesarethe courtesy of the NationalLibrary of Finland.

Ifwesupposethatthe sixparts of eachjournalvolume werepublished atregularintervals,then the publishing dateof RevueBryologique 17(3)mightbe May-June. However,the deathofthe HungarianProf. Ch. Demeteron 12th Marchwasannounced therein. Based on this,the dating of part 3would be:1890 (post 12Mar). Toconfirmthe exactpublishing date,wechecked 10referring journalsof thosetimes.Manyof them do notmention the volume orjournalat all. Some otherjournalspublished arevieworlist of contents,but on ratherlate dates,from September-December.However, J.Bot.(Morot) 4(suppl. Bull. Bibliogr.11):XLIV, published on 1st of June 1890,mentions Rev.Bryol. 17(3) and lists the articlesbydifferentauthors,including “T.Husnot.LesPhilonotis dioïques”. Accordingly,the ratherexactdateof RevueBryologique 17(3),and hence Philonotisarnellii Husn.,is12Mar-1 Jun1890,orin practiceApr-Mai1890. Husnot’sprotologue(1890a)ismoreaccuratethanthe description in “ MuscologiaGallica”and explainswhyhe described Philonotisarnellii.One reason isthatHusnotthoughtthatLindberg’s(1867)description wasnot sufficient.Lindbergonlymentioned thathe hadone fruiting specimen and afew male ,and he did notdescribethem. Husnot(1890a)cited alsoHartman’s (1871) florabut did notgivethe completecitation of the text whichwastranslated intoFrenchforhim byM[onsieur]Kindberg(K.Kindberg). Husnotonlyrepeated thatthe perigonialleavesof P.capillaris areovate,acuminatewithastrong costa and subsquarrose,and thatthe authorwasnotsurewhetheritwasaspeciesin its ownrightorratheragracile formof P.marchica.The authorconcerned in this context isS.O.Lindberghimself,who helped in writing the moss part of Hartman’sflora(see Koponen &Isoviita, 2005).

Husnot’sletters toS.O.Lindbergand V.F.Brotherus Inthe archivesof the FinnishNationalLibrary arefour of T.Husnot’s letters toS.O.Lindbergand seven of hisletters toV.F.Brotherus (see Koponen &Isoviita, 2005;Koponen &Piippo,2002). Theseletters explain whatactually happened. 80T.Koponen &P.Isoviita

Lindberg(1879) published the combination Philonotisfontana var. capillaris in his“ MusciScandinavici ”. The workwasreviewed for“ Revue Bryologique ”byVenturi(1880). Husnotclearlyreceived aletterfrom S.O.Lindberg,and answered (originalFrench,Fig. 1): Cahan,January 14,1885 Isend toyouto-dayin mail the volume 1884 of Revue,sinceyouhavereceived severalnumbers in badshape. Icould begin publishing in number2the manuscript,whichyouwill mail tome (number1will bepublished after8days). (Note1) Iwill probablyfind toyoubryologists,who wanttobuy bryophytesfrom you; and howtosend the moneytoyou?InFrancedoesnotexist,asfarasIknow,postal ordertoRussia(suchexist toSweden). (Note2) Tomymind 42cents isnottoo expensiveforrarebryophytes,whicharecoming from awell-knownbryologist. Ifyoucansend tomespecimensaccording toenclosed list,youtell me howto forwardthe moneytoyou.(Note3) Ido nothave Amblystegiumleptophyllum. T.Husnot

(Note1)LindbergsenttoHusnotamanuscriptdated 17.9.1885 (Lindberg1886a). However, the manuscriptmentioned in Husnot’sletterwasprobablyalongerone,“Sur lamorphologie desmousses”,divided between three numbers of “Revue” (Lindberg1886b). (Note2)S.O.Lindbergheld alarge duplicatecollection of hepaticscollected byhimself and hisstudents and collaborators.The aim wastocontinuepublishing the exsiccatecollection “HepaticaeScandinavicae” (Lindberg&Lackström 1874). Arnell (1884a)stated in his reviewof the exciccate:“AsProfessorLindberghasrichmaterialsforfollowing fasciculi in store,itistobehoped thatthe continuation will soon appear.” However,thisprojectdid not materializeduring Lindberg’slifetime. The collection wasdistributed morethanacentury afterhisdeath(Piippo 1993-1997). Beforeachieving independencein1917,Finland wasa Grand Duchyof ImperialRussia. (Note3)Inthe enclosed list,“Muscidesiderati” (Fig. 2),45 mossesareenumerated. Most of them arespeciesdescribed ordealtwithbyLindberg,and cited byhim in “Musci Scandinavici” orreported byArnell (1882,1884a, b). Philonotiscapillaris and P.parvula are on Husnot’slist.Lindbergevidentlydid notmail the specimensrequested.

During most of hislife Lindbergwasin poorhealthand in hislast years he worked keenlyon anumbercollections,suchasthosefrom Siberia(Koponen &Isoviita, 2005). Hedied on 20 February 1889. Atthattime Husnotwas completing the first volume of “ MuscologiaGallica”. Hewascompletelyconfused byLindberg’s(1879) treatmentof Philonotiscapillaris asavariety of P.fontana . The innerpergonialleavesof P.fontana areshort acutetoobtuseand the costa isweak,while thoseof P.marchica arelong acuminate,±erectwiththe costa percurrent.Husnothadnotyetseen the originalspecimensof Lindberg’s P.capillaris. HesentalettertoV.F.Brotherus (Fig. 3): Cahan,parAthis(Orne) 9October,1889 DearSir, Iaskfrom youif youcangivemeaspecimen from Lindberg’s Philonotiscapillaris withmale leaves( ( ). Ihaveone M.Arnell’sspecimen,but itdoesnotcorrespond towhatM.Lindberg,who hasitonlyavariety of P.fontana ,issaying. M.Arnell’s P.capillaris hasvery long and very acuminatemale leaves. Very trulyyours T.Husnot Philonotiscapillaris Lindb.and P.arnellii Husn.;one moss,twonames81

Fig. 2.Appendix“Muscidesiderati” of P.T.Husnot’slettertoS.O.Lindberg. 82T.Koponen &P.Isoviita

Fig. 3.P.T.Husnot’slettertoV.F.Brotherus.Originalinthe Archivesof the NationalLibrary of Finland. Philonotiscapillaris Lindb.and P.arnellii Husn.;one moss,twonames83

Brotherus answered thatthe herbariumwasnotavailable (see Husnot, 1890a). The reason wasprobablythatLindbergkepthisherbariuminthree differentcollections:Scandinavian,Europeanand extra-European,and Brotherus could notfind the Danishand Swedishspecimens.Instead,Brotherus sentto Husnotaspecimen collected byR.Spruceinthe Pyrenees(see the citation below) and identified byLindbergas Philonotiscapillaris .Itmaybeadded thatV.F. Brotherus neverbelonged tothe staff of the BotanicalMuseumofthe University of Helsinki,but worked athome (Koponen,1984;Koponen &Piippo,2002). Husnot(1890a)explained thattosolvethe problem of Lindberg’s Philonotiscapillaris, he hadonlythe Sprucespecimen identified byLindbergand hisnoteinMusciScandinavici(1879). Husnotfound the Sprucespecimen tobe differentfrom the Swedishspecimenssenttohim byH.W.Arnell,and he wrote on P.arnellii (1890b):

“ P.Arnellii. P.fontana var. capillaris Arnell” “--- Jen’aivu quedesexemplairessuédoisde cettevariété;elle est trèsdistinctede l’échantillon de P.capillaris desPyrénéesdeterminé parLindberglui-même et qu’il considèrecomme une variétédu P.fontana ,je ne puisdoncl’appeler capillaris etje luidonne le nom mon ami Arnell quim’en aenvoyédebeaux exemplaires.”

H.PHILIBERT’SSTUDIES

Philonotiscapillaris and P.arnellii Philibert (1894) discussed Philonotiscapillaris and P.arnellii atsome length , and came tothe conclusion thattheyaredifferenttaxa.Healsodescribed athirdspecies, P.ryanii, in thisgroupofmosses. LaterPhilibert (1897a),afterreceiving morespecimens,returned tothe topic.HethoughtthataFrenchspecimen from Valsin Ardèche represented a typicalplantof Philonotiscapillaris (T.Jensen collection from Denmark),and found thatthe otherFrenchspecimensfrom Normandyand Brittany,aswell as Belgianand Germanpopulationsof P.capillaris, wererathersimilar.Some Scandinavianplants werealittle different.Philibert (1897a)solved the problem bygiving differentpopulationsvarietalnames.The plantfrom Ardèche and other Frenchspecimenswerenamed var. gallica and aspecimen from Sweden was named afterits collectorasvar. thedenii.Philibert hadalsoreceived more specimenscollected byRyanin1895 and found them rathersimilarto P.capillaris. Thesewerenamed P.capillaris var. norwegica,and the T.Jensen specimen var. danica. Philibert’s(1894,1897a)problem seemstobethe same one Husnothad: he hadnotseen Lindberg’soriginalmaterial. Nevertheless,the originalspecimens of Philonotiscapillaris and P.parvula hadnotbeen lost.H.Philibert asked for them on loanand published (1897b)anarticle “Les Philonotis de l’herbierde Lindberg”. The materialwassenttohim byHarald Lindberg,the son of S.O.Lindberg. The specimensstudied byPhilibert (1897b)weretwosterile syntypesof P.capillaris (Nacka, 1861;Huddinge,1863; see below). The third specimen contained male plants and wascollected on 21June 1874by S.O.Lindbergin“Fennia, Ladoga, Kirpadalaks[sic, =Kirjavalaks], in fissurislatis 84 T.Koponen &P.Isoviita

[sic, =satis]humosismontisKolanaki [sic, =Kotomäki], 21. junii 1874” and named byLindberg“Philonotiscapillaris !! =var. Philonotisfontanæ ”. Twoother specimenshadsporophytes:the T.Jensen collection from Denmark(=syntype, see below)and one of the specimenscollected byI.Hagen in Norway “Smålenene,TorpiBorge” in 1887(H-SOL;Sexherb.H.Lindberg,one of the syntypesof P.ryanii). Philibert’sfinalconclusion wasthatthe specimensfrom France,Sweden,Norway,Denmarkand Finland belong tothe same species, P.capillaris, exceptfor P.arnellii, whichoccurs in alimited areain Sweden. Inthis wayhe informallysynonymized P.marchica var .tenuis Boulay, P.boulayi Corb., and P.ryanii H.Philib.

Philonotisparvula Lindb. ex H.Philib.

Lindberghimself wasnotquitesureofthe identity of Philonotisparvula. Most of the specimensnamed in hisherbariumas“ Philonotisfontana var.” are P.fontanas.str. Twospecimensnamed byhim “var. parvula ”are P.capillaris ,and one is P.tomentella Mol. (see the list of synonymybelow). Afourthspecimen, “ Philonotisfontana !formaadvar. parvulam Lindb.–[Finland.]Lojo,Skraatila, adrup. fiss.,2.VIII.1878S.O.L.”,is P.fontanas.str. Philibert (1897b)hadthese four specimensathand and discussed them atlength,finallydescribing P.parvula and P.parvula var. bomanssonii.

THE USE OF THE NAMES PHILONOTIS CAPILLARIS AND P.ARNELLII

Philonotiscapillaris and P.arnellii havebeen accepted differentlyin differentareasand atdifferenttimes.IncontinentalEuropeanliteraturethe name P.arnellii hasbeen most frequentlyused. Limpricht(1893)seemstohavebeen the first toaccept P.arnellii and,atthe same time,Lindberg’s(1879) ownsolution atthe varietallevel as P.fontana var. capillaris. Accordingly,Limprichthasthe same taxon asaspeciesand asavariety of anotherspecies.Bryhn (1899) accepted P.capillaris , P.arnellii and P.ryanii and described afourthtaxon in the group, P.media .Incommon withafewotherbryologists,Paris(1896,1905) cited P.capillaris from Lindberg(1867)and accepted Husnot’s P.arnellii asaseparate species.Severalearlywriters (e.g. Roth,1904;Warnstorf,1905) aswell asagreat numberof morerecentflorasand checklists (e.g. Podp{ ra, 1954;Pilous &Duda, 1960; Gams,1973; Corley etal.,1982; Frahm &Frey,1983; Ochyra etal.,2003) called thismoss P.arnellii.Some newflorasand checklists (Casas etal.,2006; Aleffi etal .,2008) refertothe latest Europeanchecklist of mosses(Hill etal., 2006)wherethe name P.arnellii isused. Toreplace Philonotiscapillaris ,Möller(1925) eventuallyintroduced the name P.arnellii intothe Fennoscandianliterature. Subsequentlyits usage has been consistent(e.g. Weimarck,1937; C.Jensen,1939;Nyholm,1960,1998; Koponen etal.,1977; Hallingbäck etal.,2006,2008). InRussianmoss florasand checklists the name Philonotisarnellii hasalsobeen used (Abramova etal.,1961; Savich-Lyubitskaya&Smirnova, 1970; Abramov&Volkova, 1998;Ignatov etal., 2006). Philonotiscapillaris Lindb.and P.arnellii Husn.;one moss,twonames85

However,Loeske (1906)hadalreadysynonymized Philonotisarnellii with P.capillaris, although maintaining the name P.arnellii,since“derName capillaris durchseine Vieldeutigkeitaufsäusserstekompromittiert ist,sonenne ichdie in Rede stehende Formenreihe PhilonotisArnellii Husn. emend.” After Dismier(1908,1910)haddefinitivelysynonymized P.arnellii Husn. and many othernameswith P.capillaris Lindb.(see below),foracentury Europeanauthors suchasRancken (1914),Brotherus (1923,1924),Mönkemeyer(1927),Szafran (1957),Landwehr&Barkman(1966),Allorge (1974),Raeymaekers (1981), Margadant&During (1982),Orbán&Vajda(1983),Touw &Rubers (1989) and Frahm (2005) continued tousethe name P.capillaris. InBritain,Braithwaite(1893,1905) treated Philonotiscapillaris respectivelyas P.fontana var. capillaris and asaspecies . Dixon &Jameson (1896) alsoaccepted Philonotiscapillaris atthe specificlevel and discussed the status of P.arnellii and P.ryanii.The taxaareseparated mainlyon the basisof the position of the perigonialleavesand theircosta.Theycame tothe conclusion that“It seemsprobable thatafterall theseformswill eventuallyhavetobereunited”. SubsequentlyBritishbryologists,e.g. Dixon &Jameson (1904),Ingham(1907), Duncan(1926),Warburg(1963)and Field (1963,1975,1976,1977,1985),used the name P.capillaris continuously.However,recentBritishfloras(Smith,1978,2004) accepted P.arnellii. InNorthAmerica the name Philonotiscapillaris hasbeen unanimously in use(Flowers,1935;Grout,1940; Crum etal.,1965,1973; Lawton,1971; Ireland etal.,1980,1987; Crum&Anderson,1981;Anderson etal.,1990; Allen, 2002).

THE NOMENCLATURE OF PHILONOTIS CAPILLARIS LINDB.

Welist belowthe nomenclatureof Philonotiscapillaris Lindb.,including the otherpossibilitiesif thatname isnotaccepted. Wedid notstudyall of the typesinvolved but trust Dismier’s(1908,1910)synonymizationsand citethem. The label information iscopied directlyand information from the protologuesis in squarebrackets [].

Philonotiscapillaris Lindb. Hedwigia 6:40.1867.—Bartramiacapillaris (Lindb.) Lindb. ex T.Jensen, Bot. Tidsskr. 2:272.1868. — Philonotismarchica var. capillaris (Lindb.) Limpr. in Cohn,Krypt.-Fl. Schlesien 1:117.1876.—P.fontana var. capillaris (Lindb.) Lindb.,MusciScand. 15. 1879. — P.fontana subsp. capillaris (Lindb.) Hérib., Mém. Acad. Sci. Clermont-Ferrand sér.2,14:284. 1899. — Lectotype (here designated):“359. Bartramia ,flosmasc?etfoem.” Denmark. “Mellem Björnsholm og Lundgårdivejgröft,skjultunderlyng og pors,iskygge. Logstor,IV.1863” T.Jensen (“Philonotiscapillaris Lindb.n.sp.”,H-SOL, Fig. 4a). — G.Raeymaekers marked the specimen in H-SOL asthe lectotype in 1980.— Syntypes :Denmark. “ Philonotiscapillaris Lindb.c.fr.,Majm.1863 lectaaJensen” (H-SOL, Fig. 4b);“Dania, Jylland,paroeciaLögstör,in ericetoarenaceo sub Myrica et Erica,interLundgårdetBjörnsholm,unametsingulamcapsulam!,Sept. 86T.Koponen &P.Isoviita

Fig. 4. Labelsof the types.–a.Lectotype of Philonotiscapillaris Lindb.–b.The tinysyntype with young sporophytesinside the lectotype cover.–c.Lectotype of P.arnellii Husn.

1863 leg. Th. Jensen” (S, exherb.S.O.Lindberg). Sweden. Stockholm,Nackain declivoarenoso,VIII.1861 S.O.Lindberg (H-SOL;fragmentin S);Huddinge in abruptoargillaceo adviamvaporariam,IX.1863 S.O.Lindberg(H-SOL); Drottningholm,foss.argil.,inter Br.annotin.,V.1865S.O.Lindberg(as Philonotis fontana var. capillaris Lindb.! ( )(H-SOL;fragmentin S). —Note1. [ Bartramiacapillaris Lindb. ex T.Jensen, Bot.Tidsskr. 2:272.1868. — Philonotis capillaris (Lindb. ex T.Jensen) Milde,Bryol. Silesiaca 242.1869.]—Note2. P.fontana var .tenuis Boulay,Musc.France1:217.1884,nom. illeg. incl. var.prior , P.marchica var. capillaris (Lindb.) Limpr.—Type:Homotypicwith P.capillaris Lindb.—Synonymized byDismier(1908). P.macounii Lesq. et James,Man. MossesN.Amer.208. 1884. — Type:Vancouver Island, Macoun. — Synonymized byDismier(1908). —Note3. Philonotiscapillaris Lindb.and P.arnellii Husn.;one moss,twonames87

Philonotisarnellii Husn., Rev.Bryol. 17:45. Apr-Mai1890; Muscol. Gall. 268. Jun- Jul1890.—Lectotype (heredesignated):Sweden. Småland,Barkeryd,Boarp,ad radicesrupium,2.VII.1884 H.Wilh. Arnell (GB ( ,isolectotype in S ( ,“vid skuggade bergsrötter”) (Fig. 4c). —Synonymized with P.capillaris byLoeske (1906),Dismier(1908,1910),Brotherus (1923),Mönkemeyer(1927),and Field (1985). —Note4. P.capillaris Lindb. sensu Husnot,Muscol. Gall. 269. 1890.—Based on:Pyrenees, Lesperou, Spruce (H-SOL, H;S, exherb.S.O.Lindberg). —Synonymized by Dismier(1908). —Note5. P.ryanii H.Philib., Rev.Bryol. 21:8. 1894. — Type:Norway.Skaare,Onsö,des rochers humidescouverts de terre,10.X.1893 Ryan (notseen). —Synonymized by Philibert (1897a)and Dismier(1908). P.boulayi Corb., Mém. Soc.Sci. Nat.Cherbourg 30:287,1897.—Homotypicwith P.marchica var .tenuis Boul. and P.capillaris Lindb.—Synonymized byDismier (1908). P.capillaris var. thedenii H.Philib., Rev.Bryol. 24:9. 1897.—Isotype:Sweden. Bohuslän,Hede soldattorpiTanumssocken,VI.1878 Hugo Thedenius (S, exherb. A.E.Jäderholm). —Synonymized with P.arnellii byMöller(1925). —Note6. P.capillaris var. gallica H.Philib., Rev.Bryol. 24:9. 1897.—Syntypes :France. Ardèche,Vals,1884 Philibert (notseen) and specimensfrom “desautreslocalités francaises”. —Note7. P.capillaris var. danica H.Philib., Rev.Bryol. 24:14. 1897,nom. inval.,incl. P.capillaris Lindb. ≡ P.capillaris var. capillaris.—Note8. P.capillaris var. norwegica H.Philib., Rev.Bryol. 24:14. 1897.—Syntypes :The type of P.ryanii (notseen);Norway,1895 Ryan (notseen);Norway.1895 Norge. Smålenene,TorpiBorge,17.V.1887 I.Hagen (H-SOL;S, exherb.H.Lindberg,ex herb.S.O.Lindberg[sic!]). —Note9. P.parvula Lindb. ex H.Philib., Rev.Bryol.24:86.1897(syn. nov.). — P.parvula Lindb., Hedwigia 6:40.1867 (nom. nud.). — Lectotype (heredesignated):Sweden. Västergötland,inter AmphidiumMougeotii in fiss.rupisdiabas.m.Hunneberg, VI.1859 S.O.Lindberg (H-SOL). — Syntypes :Sweden,Stockholm,arenanuda abruptaadPetersberg,10.VII.1869 S.O.Lindberg (H-SOL as P.fontana var. parvula Lindb;S);Finland. Ladoga.ins.Puutsalo,in fissumrup. 30.VI.1874 S.O.Lindberg (H-SOL as P.parvula !Lindb.). —Note10. P.parvula var. bomanssonii H.Philib., Rev.Bryol. 24:86.1897[& H.Philib.in Bomansson, ActaSoc.FaunaFl. Fenn. 18(4):97.1900.]— Type:Finland. Ahvenanmaa (Åland). [Sund,Jussböle,fuktig jordåen klippavid Bromanstorp, 15.VIII.1865]J.O.Bomansson (notseen). P.vancouveriensis Kindb.,Eur.N.Amer.Bryin. II: 326.1898. — Lectotype (here designated):N.Amer.,VancouverIsland,wetrocks26.4.1887 J.Macoun192 (S, ex herb.N.C.Kindberg). — Syntype:Alaska, Nagai-ön,18.IX.1892 J.M.Macoun 132 (S, exherb.N.C.Kindberg). —Synonymized byDismier(1910). —Note11. P.media Bryhn, Kongel. Norske Vidensk. Selsk. Skr. 1899(3):39. 1899. — Syntypes :Norway.Sätersdalen,Frøisnäs,iBygland sn[admurumviæterra obtectum], 24.VII.1894 N.Bryhn (S);Prope prædiumOse,vallisSætersdalen,Lat. sept.59°,Alt.supramare300 m,[addecliviapublicarenacea],VII.1895 N.Bryhn (S, MusciNorvegiciexherb.N.Bryhn,original!);Nedenaesamt,Kjöndalen ad terramsilvaticam,8.VIII.1891 I.Hagen (S). —Synonymised byDismier(1908).

Note1. The syntypesof Philonotiscapillaris werecollected before1867 and the lectotype must beselected from them. One T.Jensen gathering of 1863 isthe only collection having sporophytes,whichwerementioned in the protologue. However, 88 T.Koponen &P.Isoviita twospecimenswithdifferentdatesarekeptinside the specimen cover.According toT.Jensen’slettertoS.O.Lindberg,dated on 2.October1863 (in the Archivesof FinnishNationalLibrary),Jensen sentseveralspecimenscollected atdifferent times.Lindbergobviouslyfirst received aspecimen collected in April 1863 without sporophytes(Fig. 4a)and latergotatinyspecimen collected from the same locality in May1863 having young sporophytes(Fig. 4b). Weselectthe largerspecimen marked as“newspecies”byLindberghimself,although ithasno sporophytes. AthirdJensen collection originating from S.O.Lindberg’sherbarium,collected in September1863 and showing morecorrectlyspelled locality names,isin S.

Note2. Thisparagraph shows the basionymand authorcitation of Philonotis capillaris if,our argumentation notwithstanding,the validity of Lindberg’s(1867) publication isnotaccepted. Inthatcase,aspecimen seen byT.Jensen should be designated asthe lectotype. Weasked forthe Jensen collectionsof P.capillaris on loanfrom C, but havenotyetreceived them. However,the Danishspecimensused forthe typification of P.capillaris Lindb.,lectotype (Fig. 4a)and twosyntypes, formquiteessentialpart of T.Jensen’s(1868) Bartramiacapillaris material. Although Milde did notciteT.Jensen (1868),underArt.33.3of the Code Milde’sname canberegarded asavalidlypublished combination based on Bartramiacapillaris (the same appliestothe subsequentlypublished infraspecific combinationsinvolved).

Note3 .Macoun(1892)gavethe detailsof the type locality:“Wetplacesnear Victoria, VancouverIsland,May9th,1875”.

Note4.Nospecimensarelisted in Husnot’s(1890a, b)description of Philonotis arnellii. According tothe protologue, P.arnellii isbased on specimenscollected by H.W.Arnell in Sweden and sentbyhim toT.Husnot.Weasked forthe type of P.arnellii on loanfrom MuséumNationald’HistoireNaturelle,Laboratoirede Cryptogamie (PC),whereHusnot’soriginalherbariumiskept,but itcould notbe located there. Fortunately,in the bryologicalliteraturethe probable type gathering hasbeen cited severaltimes.Limpricht(1893)cited one specimen:“Wurde von Dr.H.Wilh. Arnell am2.Juli 1884 amFussederFelsen bei Boarp,Barkeryd Smaland in Schweden entdecktund als Philonotiscapillaris Lindb.vertheilt.” Roth(1904) cited the same Philonotis arnellii specimen,but gavethe wrong date“am2.Juni 1884”. Möller(1925) confirmed thatthe specimens H.W.Arnell senttoT.Husnotweremostlycollected in 1884 in Småland at Boarp,in Barkerydparish. Wehaveseen three Arnell specimenscollected in that areain 1884. Husnot’slettertoV.F.Brotherus tellsusthatHusnothadonlyone Arnell specimen withmale plants,whichHusnotdescribed. The preferred type on the basisof the protoloquewould besuchaspecimen. The specimen cited by Limprichthasonlymale plants,and,therefore,isselected asthe lectotype (Fig. 4b). The otherspecimenscollected in 1884 aresyntypes.

Note5. Itisconfusing thatHusnot(1890b)in“MuscologiaGallica”alsohas Philonotiscapillaris Lindb.Asexplained above,he hadnotseen the original specimensof P.capillaris Lindb.and V.F.Brotherus senthim adifferent specimen. AsLindberg’soriginaltype wasnotdefinitelyexcluded,Husnot’s Philonotiscapillaris cannotbetreated asalaterhomonym(cf. Art.48.1);and, moreover,its materialisconspecificwith P.capillaris Lindb.Husnot(1890b)also cited P.marchica var. tenuis Boulayasasynonym. The Sprucespecimen consists of very slenderplants withshort,notlong acuminate,leaveswithaweakercostathan Philonotiscapillaris Lindb.and P.arnellii Husn.;one moss,twonames89 isfound in typical P.capillaris .Thisvariety wassynonymized directlywith P.capillaris Lindb.byDismier(1908).

Note6. Philibert (1897a)statesthathe gotthe specimen of Philonotiscapillaris var. thedenii from “M.Jaderholm”. Onthe label of the isotype,“ Philonotis capillaris var. Thedenii Philib.Rev.Bryol. 1897pag. 9” iscited.

Note7. Philonotiscapillaris var. gallica isnotlisted in IndexMuscorum(Wijk etal ., 1967,1969) norin subsequentlists of moss names(Crosby etal.,1992etc.).

Note8. Philonotiscapillaris var. danica isnotlisted in IndexMuscorum(Wijk et al.,1967,1969).

Note9. Philonotiscapillaris var. norwegica isnotlisted in IndexMuscorum(Wijk etal.,1967,1969) norin subsequentlists of moss names(Crosby etal.,1992etc.). — Hagen hadnamed hisspecimen as P.marchica, but S.O.Lindbergidentified itas P.capillaris .Philibert (1897a)mentioned the specimen and Lindberg’sopinion. Onthe H-SOL specimen probablyH.Lindbergadded:“isaccording toPhilibert Ph. Ryani Philib.inlitt.18.10.97”(originalSwedish).

Note10 .The name Philonotisparvula Lindb. nom. nud. (1867)hadappeared in the literaturemanytimes(e.g. Hartman,1871;Schimper,1876; Zetterstedt,1876; Lindberg,1879;Venturi,1882; Bomansson &Brotherus,1894). The specimen of P.parvula collected on 10.VII.1869waslisted under P.capillaris byHartman (1871). Norrlin’s(1878) P.parvula from Lake Ladogais P.tomentella .

Note11 .The lectotype of Philonotisvancouveriensis wasidentified as P.tomentella byL.Loeske and H.Möller(in 1923). The plants havesmoothlowerleafcellsand distalleafcellswithpapillaeatthe uppercell ends.The serration of the leaf marginscomprisessingle teethorelsetheyarepartiallybiserrate. The double mamillatemarginalteethpresentin the speciesof section Philonotis arelacking. The syntype from Alaskais P.tomentella ,alsosoidentified byH.Möllerin 1923. W.M.Zalesidentified the specimen as P.fontana var. fontana in 1973.

Identification of Philonotiscapillaris The characteristicsbywhich Philonotiscapillaris hastraditionallybeen separated from otherspeciesof Philonotis arethe small size,shape and position of the perigonialleaves,narrowvegetativeleavesgraduallytapering tofiliform apex,the serration of the leafmargin and the position of the papillaeor mammillaeonleafcells.Ithasmostlybeen compared and mixed with P.marchica, P.caespitosa and P.tomentella. The lattertwotaxahavethe papillaeormammillaeofthe leafcellsatthe proximalcell end and usuallyhave double mammillateteethonthe lowerleafborder(see Koponen 2003),while in P.capillaris the lowerleafmargin issmoothand the margin in the mid toupper leafisserrateorindistinctlybiserrate. The papillosity in P.capillaris isvariable. Inmost specimensstudied the leafcellsin the lowerhalf of the leafaresmooth and the cellsof the narrowleafapexhavedistinctpapillaeatthe distalcell end orelsethe distalendsarebulging. However,in some specimens(e.g. in the lectotype of P.vancouveriensis )the leafcellsin the lowerhalf of the leafhave distinctpapillae,whichmaybeatthe proximalcell end orelsecentralonthe cell. 90T.Koponen &P.Isoviita

Inthe standardfloras(Crum&Anderson,1981;Nyholm,1998;Smith, 2004) Philonotiscapillaris isseparated from P.marchica byits smallersize. P.capillaris is0,5-1,0cmtall withleaves0,5-1.1 mm long; P.marchica is1-5 cm tall with1,3-2,3mm long leaves.The cell shape atthe leafapexisoblong or rectangular,2-4:1in P.capillaris ,while the cellsin P.marchica arelinear.The most reliable diagnosticcharacterseemstobethatonlythe distalleafcellsare papillosein P.capillaris (see abovehowever),while cellsarelinear and distinctlymammillosethroughout oralmost throughout the leafin P.marchica.

Selection of illustrations Möller1925:11-15,figs.4-10,as Philonotisarnellii;Landwehr& Barkman,1966:360,fig. 311;Lawton,1971:Pl. 115:12-16; Smith,1978:460, figs.10-13,as P.arnellii;Crum&Anderson,1981;652,fig. 309A-C; Raeymaekers,1981:22,figs.11-14;Nyholm,1998:259,fig. 217B, as P.arnellii; Smith,2004:648,figs.10-13,as P.arnellii.

Selection of specimensstudied Inaddition tothe type specimenscited above,weherelist some Philonotiscapillaris specimenswhichhavebeen discussed in the literature. Sweden.Småland. BoarpiBerghagen. Vid bergsrötter,25.VII.1884 H.W.Arnell (S ( , & , UPS ( , & );Boarpsbergetvid Karlstorp. Ien fuktig skrefva, 16.VIII.1884 H.W.Arnell (UPS ( ). —Thesemightbeamong specimenssentbyArnell toHusnotand possible syntypes,but thereisno evidenceofthat.Inaddition,therearethree specimenscollected by Arnell in the same areain 1885 (H, S, UPS), Sweden.“Vg. Hunneberg,ofvanNygård,rarissime” 22.VII.1875J.E.Zetterstedt.— Zetterstedt(1876)gavethe collection as P.capillaris and considered itidenticalto P.parvula, whichS.O.Lindberghadcollected and published (in Hartman,1871) from the same locality.Husnot(1890a)referred toZetterstedt’sopinion. Germany .Rhöngebirge,aufsandigen Waldboden derStellberge bei Bocklet,14.VIII.1871 G.Geheeb (S). Esterfeld (Vorder-Rhön )anWaldweg im Hisselswald (Sand),13.X.1870 G.Geheeb (S, exherb.S.O.Lindberg). —Geheeb(1878) came toconclusion thathis specimensand manyothers representavariety of P.marchica. France .Sur unrocherhumide,prèslagaredeBerjou-Cahan(Orne),Husnot,MusciGalliæ 733 (as Philonotismarchica Brid. Var. tenuis Boul.) (S).

NOMENCLATURAL RECOMMENDATION

The reason forusing the name Philonotisarnellii wasthatthe epithet capillaris wasconsidered confusing,having been used in differentsenses(e.g. Loeske,1906). Yet,if the type method isapplied suchold usagesarenolonger importantand,furthermore,morphotypesof asingle speciesareinvolved in this case. Wedonotconsideritreasonable underthe currentCode (McNeill etal., 2006:Arts.14and 56)that P.arnellii should beproposed forconservation against P.capillaris, northatthe lattername could beformallyrejected. Therewould be no logicin eitherof theseactionsand theyareunrealisticsince,duetothe clearly increasing role of P.capillaris during past twoyears in particular,the competing Philonotiscapillaris Lindb.and P.arnellii Husn.;one moss,twonames91 namesareabout equallyused in the literature. As P.capillaris Lindb.isthe correctname forthisspeciesunderthe Code,weabsolutelyrecommend its use. Acknowledgements.Wethank the Curators of the herbaria(C, G, PC, S, UPS) who forwarded specimenson loanormade attempts tofind type materialforour study, Mrs.MarttiinaJaatinen-Genevet,MrDaniel Genevetand Miss JuliaGenevetfor translating Husnot’sFrenchtexts forus,and the librarianatH, SirkkaSällinen,aswell as Dr.David Long (Edinburgh) forrareliterature. Weadditionallythank ProfessorTeuvo Ahtiforencouraging agreementwithour opinionsand Dr.Neil Bell forlinguisticrevision of one of the last versionsof the manuscript.

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