5hl a review of lejeuneaceae (marchantiophyta) in the russian...

Botanica Pacifica. A journal of plant science and conservation. 2019. 8(2): 85–106DOI: 10.17581/bp.2019.08208

85©Botanical Garden-Institute FEB RAS. 2019

A B S T R A C TThe Russian Far East is on the northern edge of the Lejeuneaceae distribution in PacificAsiaandthemajorityof taxaknownherearerestrictedtothesouthernmostflankof theregion.ElevenspeciesareconfirmedfortheRussianFarEast:1 – Acrolejeunea, 1 – Cheilolejeunea, 4 – Lejeunea, 4 – Cololejeunea and 1 – Microlejeunea.Allof themarereviewedinthepresentaccount,withdataondistributionwithintheRussianFarEast,morphologicaldescriptionsandfiguresbasedonmaterialscollectedintheFarEast.Inaddition,identificationkeystogeneraandspeciesareprovided.K e y w o r d s : Lejeuneaceae, Acrolejeunea, Cheilolejeunea, Cololejeunea, Lejeunea, Microlejeunea, theRussianFarEast,EastAsia,North-EastAsia,taxonomy,distribution

Р Е З Ю М ЕБакалин В.А. Обзор семейства Lejeuneaceae на российском Даль-нем Востоке. Представители Lejeuneaceae обычны только в южной части российского Дальнего Востока, севернее встречается единственныйвид –Lejeunea alaskana. ВсегонаДальнемВостоке выявлено 11 видов: 1 –Acrolejeunea, 1 – Cheilolejeunea, 4 – Lejeunea, 4 – Cololejeuneaи1–Microlejeunea.Всеониобсуждаютсяврамкахнастоящейстатьи,описывающейихраспространение,экологиюиморфологиюпообразцамсроссийскогоДальнегоВостока.Приводятсяиллюстрациивсехизвестныхврегионетаксоновиключидляопределенияродовивидовсемейства..Ключевые слова: Lejeuneaceae, Acrolejeunea, Cheilolejeunea, Cololejeunea, Lejeunea, Microlejeunea,российскийДальнийВосток,ВосточнаяАзия,таксономия,распространение

VadimA.Bakalin e-mail:[email protected]

BotanicalGarden-InstituteFEBRASVladivostok,Russia

Manuscriptreceived:25.07.2019 Reviewcompleted:30.09.2019 Acceptedforpublication:17.10.2019 Publishedonline:31.10.2019

VadimA.Bakalin

A review of Lejeuneaceae (Marchantiophyta) in the Russian Far East

Lejeuneaceaeare the largest familyamong thehepaticsacounting74acceptedgenera(Zhuetal.2019)andca1300recognizedspecies.Lejeuneaceaearecharacterizedbytermi-nal-lateral branching (Lejeunea type, rarerFrullania type, in-novationsusuallyof theRadulatype,thepresenceof asinglearchegoniumineachgynoecium(Frey&Stech2009,Grad-stein2013).Theleavesconsistof threeparts:alargedorsal,incubouslyorientedandinsertedlobe,aventralsmallerlobe,calledlobule,thatisconnectedtothelobe,andastylusthatiscommonlyvestigialorabsent.AscommentedbyFrey&Stech(2009:96),“morethan75%of theliverwortsof tro-picallowlandforestsandalmostallepiphyllsbelongtoLejeu-neaceae”.Theextraordinaryspeciationinthisgroupispro-bably connectedwith theCretaceous and post-Cretaceousexplosion of Angiosperms that provides new ecologicalniches in tropical forests (Wilson et al. 2007).Thehighestdiversity (still poorlyunderstood)occurs inhumid tropicalforests,withmoderncentersof speciation inSoutheasternAsiaandNorthernSouthAmerica(Frey&Stech2009).

Althoughtheamphi-PacificAsianregionhousesahighnumberof Lejeuneaceaetaxa(morethan500species), theRussian Far East has a very limited number of recordedspecies,andallof themarerestrictedtothesouthernflankof thearea(withoneexceptionforLejeunea alaskana).DuetotheirpeculiarappearanceamongotherRussianFarEastliverworts,confusionwithotherfamiliesseemstobehardly

possible,with the exceptionof Nipponolejeunea (Jubulaceae)which,however,hasciliateleaf margins–afeatureveryrareinLejeuneaceaeworldwide.Lejeuneaceaedidnot attractedspecialattentionof hepaticologistsintheRussianFarEast.Theearlyrecordof Acrolejeunea sandvicensis(underBrachio lejeunea)wascomprehensivelydiscussedbyZerov(1965),someotherswereonlylisted(e.g.Horikawa1940).WellknowninRussia,Schljakov’smanual(1982)forthehepaticsof thenor-thernUSSRonlyincludedLejeunea cavifolia(Ehrh.)Lindb.(ataxonabsentintheRussianFarEastaswellasinthewholeof thewesternamphi-Pacific).Withinthelastdecadesseve-ralnewrecordsof this familywerepublished.Gambaryan(1992) listedfourtaxaof thefamily,withL. cavifolia in correctlyidentified(specimensbelongtoL. japonica).Potemkin(2003)recordedCololejeunea japonicaforthefirsttimeinRus-sia. Bakalin (2007a, 2009) reviewedCololejeunea in Rus sian Asia (later onemore taxonwas found in the Russian FarEast) andCheilolejeunea in theRussianFarEast. Bakalin&Borovichev(2014)andBakalinetal.(2012)recordedLejeunea neelgherriana and L. alaskana as new for Russia. Some taxawere earlier reportedwithout descriptions for the RussianFarEast (e.g.Microlejeunea punctiformis, under Lejeunea ulicina (Taylor)Gottsche,Lindenb.&NeesinBakalin2007b).More-over, many new distributional data were collected withinlast several years. The latter circumstance, as well as thedifficultiesinidentificationof ‘lejeuneaceous’materialsfrom

86 Botanica Pacifica. A journal of plant science and conservation. 2019. 8(2): 85–106

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theRussianFarEastintheabsenceof keyshaveinitiatedthepresentwork.Itisworthmentioningthattheprominentpa-perdescribingJapaneseLejeuneaceaebyMizutani(1961)canbeusedtoidentifyalmostallLejeuneaceaeintheRussianFarEastalthough it isdifficult tousedue toenormouslyhighdiversityof thefamilyinJapanthatmakestheidentificationkeysrelativelydifficult.Themaingoalsof thepresentaccountweretoprovide:1)newdataondistributionandecologyof LejeuneaceaeintheRussianFarEast,2)identificationkeystoalltaxaknownintheRussianFarEastand3)morphologicaldescriptionsandfiguresbasedonavailablematerialfromtheRussianFarEast.ThepresentworkcontinuestheliverwortrevisionsfortheRussianFarEast(Bakalin2016,2018,2019;Bakalin&Klimova2019,Borovichev&Bakalin2016,Boro-vichevetal.2014,2015,Mamontovetal.2015,etc.)towardtothepreparationof Liverwortfloraof theRussianFarEastthatweplantopublishinthenextcoupleof years.

M A T E R I A L A N D M E T H O D SThebackgroundforthepresentworkwastheliverwort

specimens collected in theRussianFarEastby the authorwithinthepast15years.Theresultsontaxonomicdiversitywerepreliminary(asitwaspossibleatthattime)summarizedindistributiondotmapsbyBakalin(2010).

Intotalover300specimensof Lejeuneaceaewerecollec-ted.AsideaforementionedmaterialtheherbariumspecimenscollectedbyS.K.GambaryanandpreviouslyhousedinVLA(nowtransferredtoVBGI)weretakenintoaccount.Alldis-tributiondatawerere-checkedinthecourseof theprepara-tionof thepresentaccountandthedistributionprovidedforeachspecies isbasedonstudiedmaterialswithrareexcep-tion, where cited. The valuable vouchers (commonly oneper locality studied) are cited at the endof thepaper andarehousedinVBGI.Thegeneraldistributionof taxawereobtainedfromBakalin(2007a,b,2008,2009),Choi(2013),Mizutani (1961, 1982, 1998), Schljakov (1982), So& Zhu(1996),Theirs(2018),Yamada&Iwatsuki(2006)andsomeotherpaperscitedinthetext.

T A X O N O M I C T R E A T M E N TImportantfeaturesusedinidentification(andtaxonomy)

of Lejeuneaceae known in the Russian Far East are thebranchingtype(Frullaniatype[branchreplacingthelobule]and Lejeuneatype[branchjustbelowleaf insertion])andlo-buleteethfeatures,suchastherecognitionof thefirstandthesecondteeth,withtheirnumerationstartingatthejunc-tionof thelobulewiththeventralmarginof thelobe.Oilbodycharacteristicsarenotsovaluableforidentificationof taxaincomparisonwith.e.g.SolenostomataceaeorLopho-ziaceaetaxaknownintheareatreated.However, insomecasescitedinthetextandkeyoilbodiescanofferadditionalinsights to identifysometaxa: thebest feature to identifyLejeunea japonicaarenumerousandhomogenousoilbodies,Cheilolejeuneaisveryeasilyidentifiedbyafewlargeoilbodiesfilling the cell lumen in the leaf lobe.Currently 5 generawith11speciesof LejeuneaceaeareknownintheRussianFarEast.Thegeneramaybekeyedasfollow(usedfeaturesareappliedtothetaxaknownintheRussianFarEast):1.Plantsbrownishtorusty-brown,robust,with15–20longi-tudinalcellrowsintheouterlayerof thestemcrosssection,

branching predominantly of Frullania type (branches of Lejeuneatypearerare),leaf lobule3–5-toothed,underleavesundivided.............................................................Acrolejeunea 1.Plantspalegreentobrightgreenandwhitish,rarelyyel-lowish,neverbrownishorrusty-brown,withlessthan6–8longitudinaloutercellrowsinthestemcrosssection,bran-ching of Lejeunea type, leaf lobule2-toothed,underleavesbilobedorabsent......................................................................22.Plantsminute,underleavesabsent,stylus(sometimesves-tigial)regularlypresentalthoughsometimeseasilycaducous...............................................................................Cololejeunea2. Plantsminute tomoderate in size, regular underleavespresent,stylusabsentorearlyandeasilycaducous..............33.Oilbodies(1–)2–4percell,brownish,nearlyfillingcelllu-men,thesecondtoothof theleaf lobulefarlargerthanthefirstone,leavesimbricate...............................Cheilolejeunea3.Oilbodiesnumerousor5–10percell(if fewer–neverfillingcell-lumenand leavesdistant), thesecond toothof theleaf lobulesmallerthanthefirstoneorabsent,leavesimbricatetodistant..................................................................44.Plantsverysmall,lessthan0.5mminwidth,leavesdistant,lobuleonlyslightlysmallerthanlobe............Microlejeunea4.Plantslarger,widerthan0.7mm,leavesmostlyimbricate,oratleastcontiguous,lobulefarsmallerthanlobe(less1/4of thelobesize)..........................................................Lejeunea

Acrolejeunea (Spruce) Schiffn.,Hepat. (Engl.-Prantl): 128,1893(=TrocholejeuneaSchiffn.Ann.Bryol.5:160.1932;Brachio lejeunea(Spruce)Schiffn.Hepat.(Engl.-Prantl):128,1893)

Thegenusincludes21speciesandisdistributedmostlypantropically, although spreading both southward andnorthwardof thetropicalzone.Theonlytaxonknowninthe Russian Far East is Acrolejeunea sandvicensisanditslocali-tiesarethenorthernmostforthegenusworldwide.BeforeitwasknowninRussiaasTrocholejeunea sandvicensisuntilWangetal.(2014)transferredittoAcrolejeuneabasingonresultsof molecular-genetic research.Wanget al. (2014:36)circum-scribed Acrolejeunea (includingTrocholejeunea) as“possessingstems with a hyalodermis, entire-margined leaves, under-leaves,bractsandbracteoles,leaf cellswithcordatetrigonesandhomogeneousoilbodies,androeciawithepistaticbractscontainingoneantheridium,andpluriplicateperianths”.AsdiscussedbyWangetal.(2016),themainfeaturesseparatingAcrolejeunea sandvicensisfrommostotherspeciesof thegenusare thepresenceof gynoecial innovationsof theFrullania typeandamoremassivesetabuiltof 4–16innerand16–32outerrowsof cells.Inaddition,vegetativebranchesof theFrullania type aremore common inA. sand vi censis than in mostothermembersof thegenusAcrolejeunea.

Acrolejeunea sandvicensis (Gottsche) Steph., Bot. Jahrb.Syst.23(1/2,3):312,1896(≡Phragmicoma sandvicensisGott-sche,Ann.Sci.Nat.Bot.(sér.4)8:344,1857;≡Trocholejeunea sandvicensis (Gottsche)Mizut.Misc.Bryol.Lichenol.2(12):169,1961)

Description1. Plants merely rigid, brownish to rustybrown, rarely green in shady places, sometimeswith dis-colored leaf margins in well-exposed sites, in loose purepatches or in themixture with other hepatics (most fre-quentlywithFrullania) andmosses,1.4–1.8mmwideand10–20(–40)mmlong.Rhizoidssparsetonumerous,inerect1Hereandbelowthedescriptions,if otherwisenotmentioned,arebasedonspecimens from theRussianFarEast and/or featuresobservedinstudiedspecimens.

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Lejeuneaceae in the Russian Far East

toobliquelyspreadingfascicles,denselyattachingplantstothesubstratum,includingotherbryophytes.Stemyellowishbrownish,freelyirregularlybranched(branchingof Frullania type,rarelywithafewLejeuneatypebranches);crosssectionof well-developedstemsca300µmindiameter,outerlayerca18–20cellrows,cellsvaryinginsize,smallerindorsalside,wherewiththickenedwalls,25–38µmindiameter,ventrallylarger,to70µmindiameterandwiththinnerwalls,trigonessmallandconcave,innerlayer7–8cellshigh,cellwallsbe-comethickerdorsally,17–38µmindiameter.Leavesimbri-cate;lobeobliquelytosubtransverselyinserted,withdorsalmarginsometimesrecurvedwhendry,whenflattenedintheslidebroadlyovatetorounded,withroundedapex,wellde-velopedleaves1.0–1.2×0.9–1.0mm;lobuleconcave(if toviewfromventralside),semicircular tosubquadrate,withouter side rounded,with3–51–2-celled teethalongmar-gin. Cells in lobe middle thin-walled, subisodiametric tooblong, 25–50× 20–40 µm, trigones largewith all sidesconvexor concave-convex-convex (cordate), intermediatethickeningscommon(inapicalpartof theleaf presentinnearlyallcells,downwardca.20–30%of cellshavesuchkindof thickenings);oilbodies15–30andmorepercell,small,ellipsoidaltofusiform,homogenous.Underleavesca300×500µm,transverselyelliptic,mostlyappressedtothestem,with recurvedmargins (sometimesvery loosely so).Autoicous.Androeciaintercalaryinthemainaxis,with2–4pairs of bracts. Perianth obovate, pluriplicate (with 8–12plicae),welldevelopedexertedfor1/3–1/2of its length,0.8–1.0 × 0.8–0.9 mm, with one Frullania type subfloralinnovation.Figures1;2A;3D. [Capsulewall2cells thick;elatersbrown,300–400×25µm,with1–2-spirals;sporesgreen, irregularly elliptic, 65–85 × 50–70 µm, minutelypapillose(Mizutani1961)].

Comment.Theplantsareverycharacteristicinappea-rance and confusion seems to be hardly possible amongregional taxa. The entire leaf lobeswith rounded apices,small3–5-toothedlobuleandentireandwideunderleavesaredistinctive.

Ecology.Neutrophilousmeso-xerophyte.Thehabitatsof the species include open to partly shaded rocks, treetrunks,branchesinevergreentodeciduousforests.Besides,Zhu & So (2001) reported its epiphyllous occurrence insouthernChina(Fujian,Guangdong,Yunnan,XizangandZhejiang).WithintheRussianFarEastthisisobligateepili-thictaxongrowingoverrocky(of basictoneutralreactionof aqueous extract) outcrops in full sun or open (rarelypartlyshaded)sites in lowaltitudes.Commonlyoccurs inpuremats ormixedwithFrullania taradakensis Steph.,Porel la vernicosa Lindb.; in the northern extremities of thearea associated with Barbilophozia barbata (Schmidel exSchreb.)Loeske,B. hatcheri(A.Evans)LoeskeandFrullania appendiculata Steph. Among our Lejeuneaceae this is themostxerophytictaxon.

Distribution.EastAsiaandHawaii (Sunetal.2018).InJapanfromtheNorthtotheSouth,althoughabsentinHokkaido(Yamada&Iwatsuki2006).CommoninKoreanPeninsula (Choi 2013). Already Herzog (1926) noted itswide spreading northward in East Asia to 38°N; the re-centresearch‘stretched’itsdistributionasfarasto54°Nin

borealzone,whereitsoccurrencepossessesarelictalnature.Elevation reported forChina ranges from 25 to 1920ma.s.l. thatprobablydependson latitude (Zhu&So2001).Inthesouthernflankof theRussianFarEastthespeciesisdistributedmuchmoreevenlyacrossadministrativesub-unitsthandootherspeciesof Lejeuneaceaeduetoitshigherdrought-toleranceof thetaxon.Altitudinaldiapasonvariesfromca340ma.s.l. inAmurskayaProvince,thesameforbothlocalitesat49and54°N,withthelatterlocalitybeingfor the taxon the northernmost worldwide), the only lo-calityinKhabarovskTerritoryisintheBadzhalMts.(600ma.s.l.),thatisfromrockyoutcropsof thelargerivervalley.InPrimorskyTerritoryitisknownfromlowelevationsto600ma.s.l.insouthernpartof theterritory(notknowninnorthernextremes).ThespeciesisnotknowninSakhalinandKurils.

Cheilolejeunea(Spruce)Steph.,Bot.Gaz.15(11):284,1890Cheilolejeuneaisthegenusof predominantlyPantropical

distribution, although widely spreading northward to thesubtropics in theAtlantic region (Azores), Tasmania andNewZealand,andtothetemperatezoneof EastAsiaandNorth America. The genus includes over one hundredextant species, is subdivided into eight sectionsbasedonmolecularanalysis (Yeetal.2015),and includesstrikinglydifferent phenotypical groups such asCheilolejeunea s. str.with relatively narrow and bilobed underleaves and theformergenusLeucolejeuneaA.Evans (= sect.Leucolejeunea)with larger, entire underleaves.The congeneric nature of thesetwoentitieswasfirstshownbyYe&Zhu(2010)andresultedindifficultiestodescribethemainunitingfeaturesof thegenusinmorphologicalterms.Afewoilbodiesthataresomewhatbrownishincolorandfillingcelllumenareonebasiccharacterof thegenus;anothercharacter,separatingCheilolejeuneafromLejeunea(withwhichconfusioninourareaseemstobereadilypossible),isthewell-developedsecondtoothandthereducedfirstone(versuswell-developedfirsttooth andusually reduced second tooth inLejeunea).Thereduction(=absence)of thefirsttoothisevidentfromthepositionof thehyalinepapilla,whichisdistaltothewell-developed tooth (= second tooth) inCheilolejeunea, whilebeingproximaltothewell-developedtooth(=firsttooth)in Lejeunea.OnlyCheilolejeunea obtusifolia–thesmallesttaxonamongEastAsianrepresentativesof thegenus–isknownin our area.Cheilolejeunea khasiana (Mitt.) N. Kitag. men-tioned (but not treated) inGambaryan (1992) is surely amistakeof unclearorigin(cf.Bakalin2009).

Cheilolejeunea obtusifolia(Steph.)S.Hatt.,J.HattoriBot.Lab. 18: 116, 1957 (≡Harpalejeunea obtusifolia Steph., Sp.Hepat.5:265,1913)

Description. Plants green to deep green, yellowishgreenandgreenishyellowish,commonlybrownishyellowin the herbarium, 400–600 µmwide and 5–15mm long.Rhizoidsvirtuallyabsent.StemwithLejeunea-typebranchesand1–2Radula-typesubfloral innovationsthatcommonlysoonbecomefertilizedagain;stemcrosssectionca70µmindiameter,with6–7rowsof outercells,20–25µmindia-meter,innercellsin6rows,7–12µmindiameter,cellwallsslightlythickenedthroughout.Leavescontiguoustoimbri-


Bakalin

cate,lobesconvex,obliquelyoriented,withapexcommonlycurvedtoventralside,whenflattenedintheslide250–320×200–250µm,obliquelyovate,withroundedapex,marginentire;lobuleconvex(if toviewfromventralside),involuteinward,nearlyovate,ca.170×150µminwelldevelopedplants, thefirst tooth reduced to1cell, the second toothprominent,1(–2)cellswideand1(–2)cellslong,withend

celllongandcurvedtothekeel,keelrounded.Cellsinleaf lobemiddle12–20×12–17µm,thin-walled,trigonesmo-derateinsize,slightlyconvex,intermediatethickeningsra-relyoccursinlongersideof thecell;oilbodies2–3percell,irregularly oblong,finely papillose, filling cell lumen.Un-derleavesregular, rounded,120–150µmindiameter,1.7–2.2of stemwidth,sinusnarrow,to2/5of underleaf length.

Figure 1 Acrolejeunea sandvicensis(Gottsche)Steph.:1–planthabit,dorsalview;2–shootfragment,ventralview,withLejeuneatypebranching;3–shootfragment,ventralview;4–shootfragment,ventralview,withFrullaniatypebranching;5–leaf lobuleapicalpart;6–stemcrosssection;7,8–underleaves;9–leaf (allfromP-68-21-11,VBGI).Scales:a–1mm,for1;b–1mm,for2–4;c–100µm,for5,6;d–1mm,for7,8;e–500µm,for9



Autoicous.Androeciaterminalorintercalaryonleadingaxisoron shortbranches.Perianth rarelyproduced,distinctly5–keeled (2 – ventral, 2 – lateral, 1 – dorsal), with keelsshallowlycrispate,exertedfor1/2–3/5of itslength,notorshortlybeakedatthemouth,ca.500×400µm,bracteoleaslargeasbract,incisedbyU-shapedsinusdescendingto1/5–1/4of the length,bractswithboth lobesof similarsize.Figures2B;3E;4.

Comment. This species is the only representative of thisgenusinRussia.ItresemblessmallLejeuneafromwhichitisdistinguishablebywell-developedsecondtoothinleaf lobule(asindicatedbythehyalinepapillaatthelobuleapexbeingdistalinpositiontothewell-developedtooth)andafewinnumberoilbodiesinleaf cells(observableinfreshmaterialonly).

Ecology. Acido- and basi-tolerant neutrophilous me-so-xerophyte.Commonly itoccupiesopenorsemi-shadedrockysubstratesof neutralreactionand(rarely)treetrunks;inJapanalsoreportedfromlimestoneanddecayingwood(Mizutani1982).Within theRussianFarEast itcommon-ly occurs in semi-shaded cliffs in coniferous (oro-hemi-boreal) forest belt, as exclusion penetrates to stony fields(where in the crevices) in themountains (1660m a.s.l. insouthernmost Primorsky Territory). Commonly formspurepatches.Itworthnoting, inthenorthernextremeof thearea(BadzhalMts.at50°N),thespecieswasfoundonlyoverrockoutcropswithscatteredDuschekia, Picea, Pinus pu

mila(Pallas)RegelandWeigela suavis(Коm.)L.H.Baileyin-termingledwith alpine tundra plots in steep slopes at thediapason between 1570 to 1640m a.s.l., where once col-lectedwith pronouncedly arctomontaneFrullania sub arc tica Vilnet,Borovich.etBakalin.Thespeciesformingpurepat-chesinthemajorityof otherareas,althoughiscommonlygrowinginIturupIslandasadmixturetootherhepaticslikeDiplophyllum taxifolium (Wahlenb.)Dumort.,Eremonotus myrio carpus (Carrington)Lindb.etKaal. exPearson,Metzgeria lind bergiiSchiffn.,Plectocolea infuscaMitt.s.str.Nearby,inShi-kotanIsland,thisspeciesformspurepatches,althoughoncewas collected inmoist cliffs togetherwithMarsupella tu bulosaSteph.andmorethanoncewithPreissia quadrata(Scop.)NeesandPlectocolea ovalifolia (Amakawa)BakalinetVilnet.

Distribution.ThisspeciesgenerallyischaracterizedbyaJapanese-Koreanoro-temperatetooro-hemiborealdistri-bution,whereknownacrossbothcountriesthenbecomingmuchrarerinsouthwardextremes.Thesouthernmostout-post of the species area (ca. 28°N) based on specimensfromZhejiang,China,wasreportedbySoandZhu(1996)thatis.Thecitedauthors(l.c.)alsosuggestedoccurrencesof thespeciesinotherprovincesof Chinalyingnorthwardof Zhejiang.Thispointof viewlookshighlyprobable,ta-kingintoaccountthedistributionof thetaxonintheRus-sianFarEastmainlandsouthernextreme,wherethespeciesissparselydistributedalthoughsometimesislocallyabun-dant.The elevations across Japan (where fromHokkaido

Figure 2Planthabits:A–Acrolejeunea sandvicensis(Gottsche)Steph.(Kh-31-11-16,VBGI);B–Cheilolejeunea obtusifolia(Prim-16-6-16,VBGI);C–Lejeunea alaskana(R.M.Schust.etSteere)InoueetSteere(Kh-17–38–16,VBGI);D–Lejeunea japonicaMitt.(P-37-16-14,VBGI);E–Lejeunea neelgherrianaGottsche(V-4-57-17,VBGI);F–Microlejeunea punctiformis(Taylor)Steph.(V-2-18-16,VBGI)


Bakalin

toKyushu) varies from400m a.s.l. inHokkaido toover1000ma.s.l.inShikokuandKyushu,withthehighestobser-vedlocalityinSaitamaPrefecture(Honshu)at1700ma.s.l.WithintheRussianFarEastCheilolejeunea obtusifoliaisknowninKhabarovskTerritory,whereinrelativelyhighelevations(1570–1640ma.s.l.),onlyinBadzhalMts.WithinPrimorskyTerritoryitisdistributedinitssouthernthird,whereeleva-tionvaries from350 to1660ma.s.l. thatcorresponds tovegetation gradient from mixed Manchurian hemiborealforesttostonyfieldsabovethesubalpinebelt.InSakhalinProvinceknownbothfromSouthKurils(Shikotan,Iturup)andsouthernpartof Sakhalin,throughoutinloweraltitudes(15–320ma.s.l.),notpassingthehemiborealforestbelt.

LejeuneaLib.,Ann.Gen.Sci.Phys.6:373,1820Description. Plants 0.7–1.3mmwide, pale to bright

greenandyellowish,sometimesglossy,inloosemats,looselyattachedtothesubstratum.Rhizoidsafew,inshortobliquelyto erect spreading fascicles from underleaf bases. Stemsfreelyregularlytoirregularlypinnatelybranched,branchingof Lejeunea-type,crosssectionwith6–10outerand12–35innercellrows,outercellslarger,ventralmerophyte2cells

wide.Leavescontiguoustoimbricate,lobeconvex,slightlytostronglycurvedtoventralside,lobulenearlyplanetoin-voluteinward,withtwoteeth,thefirsttoothlargerthanthesecond,slimepapillanearproximalbaseof thefirsttooth(thusbetweenthefirstandthesecondteeth).Cellsinlobemiddlewiththintosomewhatthickenedwalls,intermediatethickenings regularlypresent (although sometimesa few),cuticlesmooth(ourtaxa),oilbodiessmallandmorethan5percell.Underleavesregular,bilobed(ourtaxa),lobesacutetoobtuse, equal ornearly so.Autoicous (our taxa).And-roeciaintercalarytovirtuallyterminalinleadingaxesorinlateralbranches.Perianthonbranches,with1Radula-typeinnovation,distinctly(4–)5-keeled,somewhatbeaked.

Oneof thelargestgenerainLejeuneaceae,countingforover170taxaandbeingmostdiverseinCentralAmerica,EastAfrica(includingadjacentislands),EastandSoutheastAsia.Inmanyareastherealdiversityremainspoorlyunder-stood(alsoduetooccurrenceof manyundescribedtaxa).Withinourflorathegenusmaybedistinguishedbyplantsmoderateinsize(0.7–1.3mmwide),ventralmerophytetwocellswide,thin-walledleaf cells,regularandbilobedunder-leaves,thefirsttoothbetterdevelopedthanthesecondone

Figure 3Planthabits:A–Cololejeunea macounii (Spruce)A.Evans (Prim-16-36-16,VBGI);B–Cololejeunea ornataA.Evans (P-38-5-16,VBGI);C–Cololejeunea subkodamaeMizut.(Prim-16-26-16,VBGI).Oilbodies:D–Acrolejeunea sandvicensis(Gottsche)Steph.(Kh-31-11-16,VBGI);E–Cheilolejeunea obtusifolia (Steph.)S.Hatt.(S-48-8-16,VBGI);F–Lejeunea alaskana (R.M.Schust.etSteere)InoueetSteere(S-18-18-17,VBGI);G–Lejeunea japonicaMitt.(P-37-16-14,VBGI);H–Lejeunea neelgherrianaGottsche(EB-5-9-15,VBGI);I–Microlejeunea pun cti formis (Taylor)Steph. (V-10-49-17,VBGI); J–Lejeunea japonicaMitt. (P-37-16-14,VBGI);K–Cololejeunea japonica (Schiffn.)Mizut.(J-1-19-13,VBGI);L–Cololejeunea macounii(Spruce)A.Evans(Prim-16-36-16,VBGI);M–Cololejeunea ornataA.Evans(P-38-5-16,VBGI);N–Cololejeunea subkodamaeMizut.(Kh-14-35-16,VBGI).Scale:50µm,forF–M



(hencehyalinepapillaatlobuleapexproximalinpositiontothebest-developedtooth),andoilbodiessmallandmorethan5percell.FourtaxaareknownintheRussianFarEastthatmaybekeyedasfollow:1.Leaf lobeswithapicesacutetoapiculate,plantscalciphi-lous....................................................Lejeunea neelgherriana1.Leaf lobeswithroundedtoobtuseapex,plantsneutro-philous.......................................................................................22.Leaf lobesstronglycurvedtoventralside,underleavesca0.5–0.7of lobeinsize,arctic-alpine.......Lejeunea alaskana2.Leaf lobesconvextonearlyplane,notorlooselycurvedtoventral side,underleaves less than1/3of lobe in size,temperate..................................................................................33.Firsttoothof lobuleprominent,2–4cellslong,2–3cellswide,oilbodies10–12percellinlobemiddle,granular..................................................................................Lejeunea otiana3.First toothof lobulesmaller,1(–2)cell longand1cellwide,oilbodiesmorenumerous(15–25andmorepercellinlobemiddle),homogeneous...................Lejeunea japonica

Lejeunea alaskana(R.M.Schust.etSteere)InoueetSteere,J.HattoriBot.Lab.44:330,1978(≡Hygrolejeunea alaskana R.M.Schust.etSteere,Bull.TorreyBot.Club85(3):190,1958)

Description. Plants prostrate to ascending, yellowishgreen to pale yellowish green, 0.7–0.7 mm wide and7–12mm long.Rhizoids virtually absent or in short fas-cicles originating from underleaf bases, nearly colorless.Stems usually sparsely pinnately branched, cross sectionslightly transversely elliptic, ca. 110–120 × 100–110 µm;outercellsin7rows,32.5–50.0×20–25μm,slightlythick-walled,yellowishgreen,innercellssmallerthancortical,20–25×10–20μm,in(12–)14–16rows,thin-walled,trigonesconcave to rarely triangular. Leaves imbricate, obliquelyspreadingandobliquelytosubhorizontallyoriented,lobesobliquely ovate to oblong, strongly convex, 650–800 ×450–550µm,marginentire,apexbroadlyrounded,curvedtoventralside;lobulesstronglyinflated,somewhatinvoluteinward,ovate-oblong,150–180×200–250µm,apexcon-stricted, first tooth subacute, indistinct, second tooth al-mostdisappearing,hyalinepapillaontheproximalsideof first tooth, keel strongly arched.Cells in the lobemiddle30–38×20–25μm,withslightlythickenedwalls,trigonesconcave, intermediate thickenings distinct to virtually ab-sent; oil bodies granulate, irregularly oblong to fusiform,4–13percell.Underleavescontiguoustoimbricate,broadly

Figure 4 Cheilolejeunea obtusifolia (Steph.) S.Hatt.: 1 – plant habit, dorsal view;2–shootfragmentwithperianth,dorsalview;3–shootfragment,ventralview;4–8–underleaves;9–stemcrosssection;10–leaf lobule,ventralview(1,3–10–fromPrim-16-3-16;2–fromPrim-16-6-16,allinVBGI).Scales:a–1mm,for1;b–500µm,for2,3;c–500µm,for4–8;d–100µm,for9,10


Bakalin

ovate to orbicular, cordate at base, 4–5 times aswide asstem,400–450×450–500µm,dividedbynarrowsinusdes-cending 1/3–2/5of underleaf length into two triangularlobes.Figures2C;3F;5:1–13.

Comment.Averyeasilydistinguishablespecies,withanunusualdistribution.Whereas thepresentspeciesshowsadistinctlyarctic-alpinerange,othertaxaof thegenusintheRussianFarEastarelimitedtotemperatecommunities,rarelyoccurringinhemiborealforests.Thepeculiarmorphologicaltraits include largeunderleaves ca. 0.5–0.7of lobe in sizeandleaf lobesstronglycurved(toalmostinvoluteinsomephases)toventralside.ThedistinctionfromtheJapanese-Korean, south-temperateLejeunea compacta (Steph.) Steph.remainsunclear(asidefromgeographicaldistribution).

Ecology. Basiphilous to neutrophilous meso-hygro-phyte.Occursinmountainmossytundras(includingedgeof solifluction spots) inopen sites, aswell as inopen topartlyshadedmoistcliff crevices(includingpurelimestoneinSakhalin!)inalpineand(rarely)subalpinebelts.Inpuremats or, in the cliffs, with admixture of hepatics, e.g. inKhabarovskTerritorytogetherwithAnastrophyllum assimile (Mitt.) Steph.,Frullania davuricaHampe exGottsche,Lin-denb.etNees,F. subarctica, Herbertus dicranus(Gottsche,Lin-denb.etNees)Trevis.,H. arcticus(InoueetSteere)Schljakov,Sphe nolobopsis pearsonii(Spruce)R.M.Schust.,andTrilophozia quinquedentata (Huds.) Bakalin. In subalpine belt (Pinus pumila thicketswithmany limestoneoutcrops in SakhalinIsland) together with Frullania subarctica, Metz ge ria pu bescens (Schrank) Raddi, Plagiochila porelloides (Torr. ex Nees)Lindenb., Schljakovianthus quadrilobus (Lindb.)Konstant. etVilnet,Trilophozia quinquedentata.Northward,toKolymaUp-landovermoisthummocksinmossytundratogetherwithAneura pinguis (L.) Dumort., Frullania subarctica, Plagiochila arc ticaBryhnetKaal.; inbasiccliff creviceswithFrullania subarctica, Mesoptychia heterocolpos (Thed. exHartm.) L. Sö-derstr. et Váňa, Radula prolifera Arnell, Scapania simmonsii BryhnetKaal.,andTrilophozia quinquedentata.

Distribution.AnArctic-alpinemega-Beringian taxon.Thedistributionispoorlyunderstood:thespecieswasdes-cribedfromArcticAlaska,whereitmaybelocallyabundant,then recorded inMagadanProvince (Bakalin et al. 2012),and later was found in several localities including KodarRange in Eastern Siberia, South-East Taimyr (the wes-ternmostknownlocality,Fedosovetal.,2015),Tardoki-YaniMts.inKhabarovskTerritory(southernmostlocalityworld-wide,knownintheareawithmanyotherarcticalpinerelicts,Bakalin 2015), after recorded in East-SakhalinMountains(alsoinrelictenvironments,unpublishedrecord).Thereportof Lejeunea cavifolia for lower courseof LenaRiver (Kon-stantinova&Filin1997)belongtoL. alaskanatoo(authorwasable toseemicrographsfromthespecimenonwhichthecitedreportisbased,kindlyprovidedbyDr.V.Fedosov,MW).The species is probably distributedwider than it isnowknown.ThroughoutitsdistributionacrosstheRussianFarEast(MagadanProvince,KhabarovskTerritory,SakhalinIsland) it does not descend below 850 m a.s.l. (in EastSakhalinMts.,whereinsubalpinebelt).Itisknownininnerareasof KolymaUpland(OlskoyePlateauandKilganskiyeMts., from1000to1400ma.s.l.), then in two localities in

KhabarovskTerritory(BadzhalMts.andTardoki-YaniMts.,from1570to1884ma.s.l.).SakhalinIsland,despiteitsmoresouthern position thanBadzhal, provides elevations from850 to 1385 m a.s.l. in one locality (East Sakhalin Mts.)only.The speciesmeetsL. japonica in the same localflorainsouthernpartof mountainousKhabarovskTerritory inTardoki-YaniRange,wheredistributionof L. japonicastopsbelow750ma.s.l.,whereastheonlylocalityobservedforL.alsakanalyingat1884ma.s.l.

Lejeunea japonicaMitt.,Trans.Linn.Soc.London,Bot.3(3):203,1891

Description.Plantsdeepgreentoyellowishgreenandgreenish, rarely yellowish, distinctly glossywhen dry, 0.8–1.3mmwide and 10–20(–30)mm long, in loose patches,commonly among or over bryophytes. Rhizoids virtuallyabsent, if present a few, colorless, separatedor inunclearfascicles,erectspreadingfromunderleaf bases.Stemsyello-wishgreenish,freelypinnatelybranched;crosssectionslight-lytransverselyelliptic(ca.100×110µm),with6–8rowsof outercells,thatarelargerthaninner,25–32µmindiameter,inner cells in 18–24 rows, 9–13 µm in diameter, trigonesvestigial throughout. Leaves contiguous to imbricate,obliquelytoveryobliquelyoriented;lobe370–600×270–480µm,obliquelyovate,withsomewhatobtusetoroundedapex;lobulevaryinginsize,fromverysmallindepauperateplantto125×110µm,withthefirsttooth1-celled,obtuse,thesecondtoothas‘crenation’nearthefirsttooth,reduced.Cells in lobemiddle subisodiametric to oblong, 25–38 ×25–30 µm, thin-walled, with small, concave to triangulartrigones, intermediate thickenings common in longer cellwall, oilbodies small, spherical to fusiform,homogenous,(15–)25–40percell.Underleavesdistinctlywiderthanlong,2.5–3.5timeswiderthanstem,somewhatcordatenearbase,appressedtothestemorobliquelytoalmosterectspreading,underleaf lobes obtuse to acute. Autoicous. Androeciaintercalarytoterminal,onshortorlongbranches,withmalebracteoles throughout the androecia. Perianth 0.7–0.8 ×0.4–0.5mm,exertedfor1/2of itslength,terminalonshortbranch,commonlywith1subfloralinnovationthatbecometo intercalaryor terminal androecia, perianthdistinctly 5–keeled(1dorsal,2lateral,2ventralkeels),keelssmoothorlooselycrenate.Figures2D;3G;6.

Comment.Therearemanyhistoricreportsof Lejeunea cavifoliafortheRussianFarEast–thespeciescloselyrelatedto L. japonica,butabsentinEastAsia.Itseemstheeastern-mostoutpostsof L. cavifoliaareintheUralandCaucasus.Eastwardof the latter (likeYakutiaandSouthSiberia,cf.Konstantinovaetal.2009)maybelongeithertoL. alaskana or L. japonica.AllspecimensseenfromtheRussianFarEastnamed asL. cavifolia really belong toL. japonica. Indeed,thesetwotaxaareverysimilarinmorphologyanddifferintheshapeof underleaveswheretheyarewiderthanlonginL. japo nicaandlongerthanwideinL. cavifolia.Besides,theunderleavesof L. japonica are cordate near the base – the featurenotobservedinL. cavifolia.Moreover,thetwotaxaarestronglydifferentindistributionwiththegapstretchingfromUralMountainstoEastAsia.Therecordof Lejeunea cavifolia for Arctic Yakutia, cited by Konstantinova et al.(2009),belongstoL. alaskana.



Figure 5 Lejeunea alaskana(R.M.Schust.etSteere)InoueetSteere:1–planthabit,ventralview;2–planthabit,dorsalview;3,4–leaf lobules,ventralview;5–8–leaves,ventralview;9–13–underleaves(1,2–fromMag-28-25-12;3–13–fromKh-19-1-16,allinVBGI).Lejeunea neelgherrianaGottsche:14–planthabit,ventralview;15–18–leaves,ventralview;19–23–underleaves;24,25–leaf lobules,ventralview;26–leaf lobeapex(allfromEB-5-9-15,VBGI).Scales:a–1mm,for1,2;b–1mm,for14;c–100µm,for3,4,24–26;d–1mm,for5–13,15–23


Bakalin

Ecology.Neutro-toacidophilousmesophyte.Thebasichabitatsforthespeciesarerocksandtreetrunks;althoughinsouthernChinaitwasreportedalsoasepiphyllous(Zhu&So2001).WithintheRussianFarEastthespeciesismalleableinecologyandpossessesitself asmeso-hygrophyticacidophyte.Itoccupiesbasesof treetrunks,stonesandrocksinpartlyshadedplaces,includingthosealongstreams,butasideof di-rectimpactof runningwater,sometimesgrowingoverdensepleurocarpousmossmatsinthebaseof boulders,rarelyonmoistfinesoilsinsteepslopesandalongstreamsinforestedareas. It occurs in broadleaved cool-temperate (multi- andmonodominant),mixed(of Manchuriantype)andrarelyoro-

hemiborealconiferousforests.Asexclusionsoncecollectedon the base of Betula lanata(Regel)V.N.Vassil.trunkinsub-alpinebeltandeveninthecreviceinstonyfieldnearthetopof OlkhovayaMt.(1600ma.s.l.),wherefoundtogetherwithFrul lania ap pen diculataSteph.andHerbertus dicranus.ThespeciesformspurepatchesorgrowingtogetherwithFrullania appendi cu lata, Lophocolea heterophylla (Schrad.) Dumort., Metzgeria pu bes cens, Plagiochila ovalifolia Mitt., Porella grandiloba Lindb.,P. vernicosa Lindb.,Radula obtusiloba Steph., Preissia quadrata overacidtoneutralreactionstonesandcliff crevices;withCo lo le jeunea ornataoverlimestonecliffs;withLophocolea minor Neesoverbarkof broadleavedtrees;withMetzgeria pubes cens

Figure 6 Lejeunea japonicaMitt.:1–planthabit,ventralview;2–planthabit,dorsalview;3–perianthousshootfragment,ventralview;4–perianthousshootfragment,dorsalview;5,6–leaf lobule,ventralview;7–stemcrosssection(1–4–fromP-61-25-08;5–7–fromPrim-19–8–16,allinVBGI).Scales:a–1mm,for1,2;b–500µm,for3,4;c–100µm,for5–7



and,asexoticvariants,withRadula japonicaGottsche,Nip ponolejeunea subalpina (Horik.) S.Hatt. over conifer trunks;withRadula complanata (L.) Dumort.,Metzgeria lindbergii Schiffn.over partly shaded Juniperus sargentii (A.Henry) Takeda exNakaibranches inSouthKurils (ShikotanIsland); inmoistcliffsonceobservedtogetherwithChiloscyphus poly an thos(L.)Corda. Once the species was collected on a wet boulderalongstream,thathadunusualassociates,includingLophozia lantratoviae Bakalin, Plagiochila ovalifolia, Scapania crassiretis BryhnandTritomaria exsecta(Schmidel)Schiffn.exLoeske.

Distribution.TemperateEastAsian,themostcommonspecies of the genus in Japan,KoreanPeninsula and thesouthern Russian Far East. In China it is reported fromGuangdong,Guizhou,Sichuan,TaiwanandZhejiang(Zhu&So2001).Thealtitudinalrangecoverselevationsfromsealeveltomiddleelevationsof mountains.Thehighestobser-vedelevationisinSichuan(2400ma.s.l.,whereitgrowsontreetrunk,cf.Zhu&So2001).WithintheRussianFarEastis known from two localities in southernpartof Khaba-rovskTerritory(765ma.s.l.,Tardoki-YaniRangeand202ma.s.l.,AnujRiverBasin),thenorthernmostat49°N.WithinPrimorskyTerritoryalllocalitieslyebetween43and44°Nandvary inelevation from low level (35ma.s.l.) to1250(ChuguevskyDistrict)and1600ma.s.l.(PartizanskyDist-rict).Within Sakhalin Province known only from severallocalitiesinShikotanIsland.TherearereportsforSikhote-AlinReserve (Gambaryan2001) and forKunashir Island(Bakalinetal.2009),wherethespecieslikelyshouldbeex-pected,butvoucherswerenotobserved.

Lejeunea neelgherrianaGottsche,Syn.Hepat.3:354,1845Description. Plants prostrate to loosely ascending at

shootapices,brightgreentoyellowishgreen,slightlygliste-ning,1.0–1.2mmwide,5–10mmlong,formingloosepat-chesovermossmats.Stemssparselybranched,80–100µmindiameter,with6–8outerand12–18innercellrows,theoutercellsthin-walled,innercellswiththickenedwallsandwithlargeconvextrigones.Rhizoidsshort,ca.150–200µmlong,colorlesstograyish,indense,erect,spreadingfasciclesoriginatinginunderleaf bases.Leavescontiguoustoimbri-cate,lobeslightlyconvextonearlyplanewhenwetandin-curved todorsal sideof shootwhendry,whenflattenedintheslideobliquelyovate,withacutetolooselyapiculateapices, 550–650× 400–500 µm; lobule 200–250× 140–160µm,inflated,slightlyinvoluteinward,withslightlycon-vexkeel,thefirsttooth1–celled,composedof triangulartowidelytriangularcell,thesecondtoothreduced.Cellsinthemiddleof the lobe20–26×15–18µm, thin-walled,withsmalltomoderatelyinsize,concavetrigones,intermediatethickenings absent,oilbodies coarselygranulate, sausage-shapedtoirregularlyfusiform,6–13×4–5µm,3–6percell.Underleavesnearlyroundedtoslightlytransverselyelliptic,350–400×350–450µm,withentiretoslightlycrispatemar-gins,dividedbyaU-toV-shapedsinusintotwolobeswithobtuseapices.Figures2E;3H;5:14–26.

Comment.Thisspecies iseasilydistinguishedamongregionaltaxabytheprominentlyacute-apiculatelobeapicesthatseparateitfromallotherknowntaxaof thegenusinourarea.

Ecology. Probably a neutrophilous mesophyte, al-thoughalsoabletogrowonacidicandbasicsubstrates.InourareaandintheadjacentKoreanPeninsula,itbehavesasacalciphilousmesophyte.KnownintheRussianFarEastfrom the only collection obtained from a limestone cliff cave(wheretogetherwithCololejeunea ornata)surroundedbycool-temperatebroadleaveddeciduousforestinsteepNE-facing slope, in area of northernmost outpost for seriesof temperate and subtropical bryophyte species (Bakalin&Borovichev2014).However,withinChinaitcommonlygrowing over tree trunks, branches and rocks, rarelyoccurringoverlivingleaves(Zhu&So2001).

Distribution.Oro-subtropicalSouthAsian(India,Ne-pal,Bhutan, SriLanka) toEastAsian taxon spreading tothewarmtemperatezone.Theoccurrencesinourareaandcentralpartof theKoreanPeninsulapossessaninterestingrelictalcharacterandareassociatedwithseveralbryophytesshowingsimilardisjunctoccurrencesamongthedominanttemperatevascularvegetation.ThenearestlocationstotheboththeKoreanandRussianoccurrencesareinsouthernJapan (Kyushu and adjacent islands, Yamada & Iwatsuki2006).Moreover,thedistributionof thistaxoninnorthernextremes(bothinRussianFarEastandKoreanPeninsula)seemstoberestrictedbylimestone,whereasinareacoreof thisspeciesitisassociatedwithtrunks,branches,rottenlogs,rarelyoccurringinrockysubstrataandlivingonleaves(Zhu&So2000).WithintheRussianFarEastitisknownfromtheonlylocality insouthernflankof PrimorskyTerritory(SestraMt.nearNakhodkaCity).ThealtitudevariesfromnearsealevelintheNorthto2520ma.s.l. inXizang(Ti-bet),whereitoccursovertreetrunks.Theanomalyindist-ributionandmorphologyof thetaxonwasdiscussedinabroadercontextbyBakalin&Borovichev(2014).

Lejeunea otianaS.Hatt.,Bot.Mag.(Tokyo)65(763/764):15,1952

Description.Plantsprostrate,yellowishgreenish(whi-tishyellowishintheherbarium),formingloosepatcheswithotherliverworts,0.8–1.1mmwide,8–15mmlong.Rhizoidsvirtuallyabsentorsolitary,colorless,erectspreading.Stemsfreelypinnatelybranched(alsocommonlyassubfloralinno-vation);crosssectionof well-developedstemsca110µmindiameter,outercellslarge,in8rows,25–40µmindiameter,innercellsin12–15rows,thin-walled,12–20µmindiameter.Leaves contiguous, lobes 600–700× 450–550 µm, ovate,withobtuseapex,slightlyconvex,withapexnotorlooselyturnedtoventralside,veryobliquelytosubhorizontallyori-ented;lobule200–220×130–150µm,ovate,withthefirsttoothprominent,acute,2–4cellshighand2–3cellswide,the second tooth reduced, 1–celled, obtuse.Cells in lobemiddlesubisodiametrictooblong,25–40×20–25µm,thin-walled, trigones small, concave, intermediate thickeningscommon. Underleaves suborbicular, mostly appressed tothestem,300–400µmindiameter,dividedbyV-toU-sha-pedsinusdescendingto1/3–2/5of thelength,lobestri-angular, merely acute. Autoicous. Perianth terminal, onleadingaxis,obovatetosubclavate,looselyfoldedinupper1/3–1/4of the lengthor inflatewith folds indistinct,ca.0.7–1.0×0.5mm,mouthdistinctlybeaked.Androeciater-minalonbranches,withorwithoutbracteoles.Figure7.


Bakalin

Comment.Arareandrelativelysmall-sizedspeciesof Lejeunea.Carefulexaminationof thelobuleteethisneededfor correct identification. Unlike the widely distributedL. ja po nica,thisspeciespossessesalargefirsttoothof 2–4cellshigh.InotherrespectsitresemblesasmallL. japonica, althoughcommonlybeingpalerandneverglossy.Somedif-ferencemaybeinoilbodiesthatarestillunknowninthepresenttaxon.

Ecology. Acidophilous mesophyte. In Japan, fromwhere itwas described, it is observedoverwet rocks in-cludingthosecoveredwithathinlayerof humusenrichedsoil (Mizutani 1961). The habitat in Shikotan Island of thesouthernKurils,wherethespecieswasonlyrecorded,isquitedistinctive.Lejeunea otianawasgathered from thin

semi-shadedbranchesindensethicketsof Juniperus sargentii thatformlargeclumpsontheisland.Thejuniperformationissomewhatof arelictnatureinShikotanwhereitformsa peculiar ‘juniper belt’ (Barkalov 2009) instead of Pinus pumila that iswidely distributed over otherKuril Islands,andistheprobableresultof aninsulareffect.If so,Lejeunea otia namayalsoberegardedas therelictof thetemperatefloradominanceintheisland.

Distribution. RegardedasarareJapaneseendemicbyMizutani (1961), it occurs throughout Honshu, ShikokuandKyushu(Yamada&Iwatsuki,2006).Lateritwasfoundinthesouthernpartof SouthKorea(Choi2013).TheonlyrecordfromtheRussianFarEastisfrom200ma.s.l.inthesouthernmostKurils(ShikotanIsland).

Figure 7 Lejeunea otianaS.Hatt.:1,3–planthabit,dorsalview;2–planthabit–ventralview;4,5–leaf lobule,ventralview;6–underleaf lobeapex;7–cellsinleaf lobemiddle;8–stemcrosssection;9–12–underleaves;13–15–leaves,ventralview(allfromK-41-16-07,VBGI).Scales:a–1mm,for1;b–1mm,for2,3;c–100µm,for4–8;d–1mm,for9–15



Microlejeunea(Spruce)Steph.,Hedwigia27(2):61,1888This isarelativelysmallgenusof Lejeuneaceaecoun-

tingca25extanttaxawiththelargestdiversityinTropicalAmericaandEastAfrica(includingadjacentislands),and,to a lesser extent, SoutheastAsia.The genus is characte-rized by distant leaves with leaf lobe commonly housedbasalocelli, relatively large leaf lobule and regularunder-leaves. Among regional taxa the confusion seems to behardly possible due to distanced leaves and small size.Following the current point of view (Söderström et al.2016),Microlejeuneaistreatedatthegenericlevelhere.

Microlejeunea punctiformis(Taylor)Steph.,Hedwigia29(2):90,1890(≡Lejeunea punctiformisTaylor,LondonJ.Bot.5:398,1846)

Description.Plantstiny,verysoft,pallid,palegreentoyellowishgreenish, in loosematsovertreebarkortogetherwithotherepiphytes,200–300µmwideand2–6mm long.Rhizoids colorless, in short obliquely spreading fascicles,‘pseudopodially’branchedatterminalends,originatedinun-derleaf bases. Stem37–50µm indiameter, freely pinnatelyto bipinnately branched (branching of Lejeunea-type), crosssection suborbicular,with 6–7outer and 3 inner cell rows.Leaves distant, lobes convex, well developed130–180 × 100–140 µm, nearly ovate, withslightlyattenuateorobtuseapex; lobuleconvex(if toviewfromventralside),welldevelopedca150–175×100µm,ovate,apexwith2teeth,thefirsttoothlarger,1–celled,ca25–30µmlong,thesecond tooth smaller,obtuse, 1–celled.Cells inleaf lobemiddle12–20×10–13µm, thin-wal-led, intermediate thickenings unclear, trigonessmallandconcave;ocelli1–2,if 2theninshortrow or separated by ordinary cells; oil bodiesgranulate, small, spherical to fusiform. Under-leavesnarrowlyobtrapezoidal,90–110×75–100µm,dividedbyV-shapedsinusdescendingto1/2of underleaf length.Dioicous.Perianthobovate(obconical),4–keeled,ca.0.4×0.3mm.Figures2F;3I;8. [Elaters120–160×12µm,colorless,indistinctlyunispiral(Mizutani1961).]

Comment.Thisspeciesiseasilyidentifieddue to its small size (even smaller than ourCololejeunea),distancedleaveswithlargelobules,regularunderleavesandpresenceof ocelliintheleaf lobebase.Thespecieswasfirstlyrecorded(Bakalin2007b)underLejeunea ulicina (Taylor)Steph., following the concept inYamada andIwatsuki (2006). However, already Zhu andSo (2001) were inclined to treat the Asianspecimensasdifferent,althoughcloselyrelatedto L. ulicina, Microlejeunea punctiformis (Lejeunea punctiformis in Zhu and So, 2001). The latterpointof viewseemsmore reasonable fromageographicalperspective.M. ulicina (T.Taylor)Steph. is restricted to easternNorthAmericaandEurope, whileM. punctiformis is anAsiantaxon.Thispointof viewwasalsoacceptedinSöderströmetal.(2016).

Ecology.Acidophilousmesophyte.Thespeciesiscom-monlygrowingontreetrunks,decayingwood,sometimeson vascular plant leaves and rarely on rocks. Withinthe Russian Far East it is restricted to epiphytic habitatsandmay be called as acidophilicmesophyte. It grows inconiferous andmixed (coniferous-broadleaved) forests inthesouthernmostKurils,thusunderaconditionof highairhumidity.OvertreebarkitformspurepatchesorgrowingtogetherwithNipponolejeunea subalpina(Horik.)S.Hatt.andNeohattoria herzogii(S.Hatt.)Kamim.

Distribution. This ismainly anEastAsian taxon stret-chingfromIndia,NepalandBhutan,thenwidelydistributedvia China (Anhui, Fujian, Guangdong, Guangxi, Guizhou,Hainan,HongKong, Jiangxi,Taiwan,Yunnan,Xizang,Zhe-jiang)andreachingKoreanPeninsulaandJapaninthenorthand South-EastAsia (Thailand,Vietnam) in the south (Zhu& So 2001).Within Japan it is distributed fromHonshu toRyukyuandBonin,butnotknowninHokkaido(Yamada&Iwatsuki 2006). Within the Korean Peninsula it is relativelycommoninsouthernflanks,butbecomesraretothemiddlepartof thepeninsula(Choi2013).WithintheRussianFarEastknownfromthesouthernmostislandsof Kurils:KunashirandShikotan,wherecollectedinlowelevations(110–150ma.s.l.).

Figure 8 Microlejeunea punctiformis (Taylor) Steph.: 1, 2 – underleaves; 3 – leaf lobule,ventralview;4–planthabit,dorsalview;6–shootfragments,ventralview(allfromK-47-26-06,VBGI).Scales:a–100µm,for1–3;b–500µm,for4,5


Bakalin

Cololejeunea(Spruce)Steph.,Hedwigia30(5):208,1891Description. Plants pale colored, greenish, yellowish

greenish to whitish, in loose mats, mostly creeping oversurface,less1000µmwideand10mmlong.Rhizoidswell-developed,inshortfasciclesbetweenlobuleadnations(twofascicles per leaf pair). Stem branching of Lejeunea-type,crosssectionwith5outerandoneinnercellrows.Leavescontiguous to imbricate, mostly strongly dorsiventrallyconduplicate, lobe nearly ovate, with rounded, acute toapiculateapex,marginentiretocrenulateduetopapillose-mamilloseprotrudences,dorsalsurfacesmoothorroughlypapillose-mamillose; lobuleca1/3of lobe insize,mostlyinvoluteinward,withtwodistinctteethof similarsize,orteethobsolete. Styli developed, small and easily caducousinolderpartstorobustandpersistent.Cellsinlobemiddlethin-walled, intermediate thickenings absent. Underleavesabsent.Autoicous(ourtaxa).Androeciaonmainaxisoronlateral branches, terminal to intercalary.Perianth terminalonbranches,5-plicate,mostlybeaked.

Cololejeunea is the largest genus of Lejeuneaceae thatcounts for over 300 extant species (somenamesmay re-presentsynonymsandmanytaxaarestillundescribed).Thegreatestdiversityoccurrsinthetropics(mostlyPaleotropicsthan Neotropics). In Japan, where Lejeuneaceae wasan object of careful revision by Mizutani (1961, etc.) itaccountsforhighdiversityof ca.50species.WithinJapanthemajorityof taxa isdistributedinitssouthernhalf (oreven fringes). Within the Russian Far East Cololejeunea occurssparselysouthwardof 48°N,beingrelativelycom-mon only in Manchurian temperate (multidominantbroadleaved or mixed) forests in the southern flanks of Primorsky Territory. Besides aforementioned, Cololejeunea areknowninthesouthernKurils,where,however,theyarerarities.Thetaxaof Cololejeuneaareeasilydifferfromothermembersof LejeunaceaeknownintheRussianFarEastbytheminute size, absenceof underleaves andpresenceof styli.FourspeciesareknownintheRussianFarEastwithone (C. japonica) belonging to subg.Pedinolejeunea Benedixex Mizut. and three others to subg. Cololejeunea. AmongtaxarecognizedinRussia,threefurtherspeciesareknown,withtwobelongingtothelattersubgenus(C. calcarea(Lib.)Schiffn. andC. rossettiana (C.Massal.) Schiffn. ) and onemore(Cololejeunea nakajimaeS.Hatt.)tosubg.Taeniolejeunea (Zwickel)Benedix.Thesystematicsatthesubgenericlevelismorphologicallybasedonfeaturesof lobemargin, leaf cuticle,positionof hyalinepapilla in leaf lobuleanddist-ributionof ocelli(absentinourtaxa).Molecular-phyloge-neticanalysisof thewholegenus,however,showedthatthesubgenera of Cololejeuneaascurrentlycircumscribedarenotwellfounded.ThetaxaknownintheRussianFarEastmaybedistinguishedbythefollowingkey:1.Papillose-mamilloseprotrudencesareabsentoverdorsalsurfaceof leaf lobe[subg.Pedinolejeunea]................................................................................................Cololejeunea japonica1. Papillose-mamillose protrudences are well developedoverdorsalsurfaceof leaf lobe[subg.Cololejeunea].............22.Terminalcellof thefirsttoothof the lobuleob-trape-zoidalandtoothsomewhathammer-shaped,apicesof pa-pillose-mamilloseprotrudencessemispherical....................................................................................Cololejeunea macounii

2. Terminal cell of the first tooth of the lobule conical,toothneverhammer-shaped,apicesof papillose-mamilloseprotrudencesconical...............................................................33. Stylus well-developed 5–12 cell long, outer surface of lobulewith papillose-mamillose protrudences throughout(with exception of a few rows adjacent to ventral lobulemargin)...................................................Cololejeunea ornata3.Stylusreduced,1–2(-3)celllong,outersurfaceof lobulewith papillose-mamillose protrudences in several rowsadjacenttothekeel......................Cololejeunea subkodamae

Cololejeunea japonica(Schiffn.)Mizut.,J.HattoriBot.Lab.24:241,1961(≡Leptocolea japonicaSchiffn.,Ann.Bryol.2:92,1929)

Description.Plantspalegreenishtowhitishgreenish,creeping,overrocksorbarkoroverotherbryophytes,700–900µmwideand5–10mmlong,stronglydorsiventrallyflat-tened.Rhizoidssparse,infasciclesor‘rosettes’frominitialareabetweenlobuleadnations,distinctly‘pseudopodiate’atthe ends. Stem 40–70 µm in diameter, sparsely pinnatelybranched,crosssectionwith5outerand1innercellrows.Leaf lobessubimbricate,whenflattenedintheslideobliquelyovate,withroundedapex,ca.40–500×300–400µm,outersurfacesmooth,marginentire;lobuleveryvaryinginshape,fromhighlyreducesandnarrowlytriangularwithapiculateapex (2-celled uniseriate end) to narrowly lingulate andtruncateintheapex,slimepapillaininnersidenearapex,thefirstandthesecondteethhardlydeveloped,welldevelopedlobes100–170×60–90µm.Cellsinleaf lobemiddle20–32×20–25µm,thin-walled,intermediatethickeningsobscure,trigones small and concave; oil bodies smooth to finelygranulate,colorlesstogreenish,5–7(–13)percell,sphericaltoellipticinprojection,3.0–3.5×3.5–5.0µm.Leavesfreelygemmiparous, gemmae distributed over inner surface of the lobe, discoid, 20–60 µm in diameter. Figures 3K; 9:1–4.[Autoicous.Maleinflorescencesinthemainaxisoronthebranchwith2–3(–7)pairsof bracts.Perianthinflated,5-keeled,0.6–0.7×0.35–0.5mm.Capsulewallhyaline,palebrown;elaters130–220×15µmwithwallssinuatelythicke-ned;sporesirregularinshape40–53×20–30µm,minutelypapillose(Mizutani1961)]

Comment.Amongregionaltaxathisspeciesisveryeasi-lyidentifiedduetosmoothleaf lobesurfacewithentirelobemargin and commonproduction of discoid gemmae overinnersurfaceof leaf lobe.ItmaybeconfusedwithpoorlydevelopedLejeunea japonica rather thanwithourotherCololejeunea,fromwhich,however,iteasilydiffersintheabsenceof underleavesandcompletelyanothershapeof leaf lobule.

Ecology.Neutrophilousmesophyte.Southwardof theRussianFarEast thecommonhabitatsof thespeciesaretreebarkandleaves(Zhu&So2001).However,inthestu-diedareathespeciesisknownfromtwocollectionsintheRussian FarEast; the first one is that byA.D. Potemkin&V.M.Kotkova (Potemkin 2003)where the specieswascollected over semi-shaded stones [of neutral reaction?]in Taxus cuspidataSiebold&Zucc.forest.Potemkin(2003)alsonotedthatthisspeciesmaybeepilithicinJapan.ThesecondRussiancollectionisalsofromcliffsinpartshadeinW-facingslopecoveredbybroadleavedforest.

Distribution.GenerallyaKorean-Japaneseoro-tempe-ratetaxon,rarelyoccurringinlowlands.Southwarditisre-



portedintwoprovincesof China(Fujian,Shanghai,cf.Zhu&Lu1995,Zhu&So2001),bothareinmiddleelevationof mountainsinoro-subtropicbelt.InJapanknownfromHonshu, Shikoku, Kyushu, Bonin (Yamada & Iwatsuki,2006), and in theKoreanPeninsula sparselyoccurs in itssouthernpart.WithintheRussianFarEast(andinRussiaasawhole)knownfromthesouthernportionof PrimorskyTerritory, in Petrova Island (altitude unknown, althoughshouldbelow)neartowesterncoastof theSeaof Japanand inMikhajklovskyRange (280ma.s.l.).Both localitiesarenear43°N.

Cololejeunea macounii (Spruce)A.Evans,Mem.TorreyBot.Club8(2):171,1902(≡Lejeunea macouniiSpruce,Bull.TorreyBot.Club17(10):259,1890)

Description.Plantsprostrate,whitishgreenishtoyel-lowishgreenish,alwayspale,inloosematsoverrocks,bark

andlargerbryophytes,commonlydenselyattachedbyrhi-zoidstothesubstratum,700–1000µmwideand3–8mmlong.Rhizoidscolorlessornearlyso,inobliquelyspreadingfascicles,originatinginareasbetweenof lobuleadnations.Stemsfreelypinnatelybranched(branchescommonlywithsmallerleaves),70–90µmindiameter.Leaf lobesimbricate,whenflattenedintheslideobliquelyovate,withapexlooselyorstronglycurvedtoventralside,dorsalsurfacewithpapil-lose-mamillose protrudences, semispherical in apices; lo-bulenearlyobovate,largeca300×200µm,withthefirsttooth 2-celled, somewhat hammer-shaped, terminal cellobovatetoobtrapezoidal,thesecondtoothtriangular,2–3cellswideand2–3cellslong,outersurfacenearlysmooth(roughly papillose-mamillose only near the keel, in apicalpartof lobule2–4cellrowsarepapillosetoo).Cellsinlobemiddle 12–17 µm, thin-walled, trigones small, concave,

Figure 9 Cololejeunea japonica (Schiffn.)Mizut.:1– lobuleandstylus,ventralview;2–gemmae;3–shootfragments,ventralview(withgemmaeproduction);4–planthabit,dorsalview.Cololejeunea subkodamaeMizut.:5–shootfragment,ventralview;6–lobuleandstylus,ventralview;7–gemma.1–4–fromKor-10-3-11;5–7–fromKh-14-35-16(allinVBGI).Scales:a–100µm,for1,2;b–500µm,for3;c–1mm,for4;d–500µm,for5;e–100µm,for6,7


Bakalin

intermediatethickeningsabsent;oilbodiessphericaltoob-long,3–6percell,veryfinelypapillose.Stylusvery small,easily caducous, 1(–3)-celled. Autoicous, freely fertilized.Androeciaonleadingaxisoronbranches,virtuallyterminal.Perianthterminalonbranches,with1subfloralinnovation,obovate, distinctly 5-keeled (2 – lateral, 2 – ventral, 1 –dorsal), roughly papillose-mamillose throughout. Capsuleshortly exerted, spherical, 200–300 µm in diameter, withcolorlesswalls.Figures3A,L;10:1–3.

Comment.Amongregionaltaxathisiseasilyidentifiedspeciesduetohemisphericalapicesof papillose-mamilloseprotrudencesandobovatetohammer-shapedfirsttoothof leaf lobule.Thisspeciesisthelargestamongregionaltaxaandmorethanotherinclinedtooccupybasiphilousrockysubstrata.

Ecology. Neutrophilous to acidotolerant mesophyte.InsouthernandeasternChinaitisknownfromtreetrunks,branches, stumps, decaying wood and leaves of vascularplantsandferns(Zhu&So2001).InJapanandtheKoreanPeninsula (the southern part of the latter) it is commonoverbarkof treesandshrubs,withrareoccurrencesoverrockysubstrates.WithintheRussianFarEastthisismostlyanepilithictaxonpreferringrocksof neutralreactionsin-cludingthosenearwatercourses(asideof directimpactof runningwater),andmaybecharacterizedasaneutrophilictoacido-tolerantmesophyte.Rarelythistaxonoccursoverbark near trunk base of coniferous trees (mostlyAbies).Cololejeunea macouniiisalmostrestrictedinourareatooro-boreal belt in themountains, although once observed instony field (above 1300 m a.s.l.) in Livadijsky Range of PrimorskyTerritory.IntheRussianFarEastitformspurepatchesoroncefoundwithC. subkodamae.

Distribution.Temperate,mostlyanEastAsianspecies,with several occurrences in Canadian British Columbia,whereitistheonlyspeciesof Cololejeunea(mainlandof Bri-tishColumbiaandMoresbyIslandof theQueenCharlotteIslands,cf.Theirs,2018).InEastAsiatheareaof C. macounii coversJapan,theKoreanPeninsula(althoughquitesparsethere),Taiwan,China(Anhui,Fujian,Guangdong,Guangxi,Guizhou,Hunan,Jiangxi,Sichuan,Yunnan,Zhejiang)andVietnam.InChinaitisknownfromaltitudesbelow1900ma.s.l.(Zhu&So2001).WithinJapanthespeciesisknownfrom Hokkaido to Kyushu, Yakushima and Ryukyu, inaltitudesthroughoutJapanlyingmostlybelow500ma.s.l.,exceedingthiselevationinShikokuandHonshu(itisworthnoting,evenintheseislandsmostof thelocalitiesof thistaxon are below 500 m a.s.l.). However, in Taiwan thisspecieswascollectedin3630ma.s.l.(Mizutani1998).WithintheRussianFarEast it occurs in oro-hemiboreal belt of mountainsinthesouthernflankof thearea.Thenorthern-most locality is in the southern extreme of KhabarovskTerritory(Tardoki-Yani),whereitwasfoundbetween540and765ma.s.l.;southwarddistributedthroughPrimorskyTerritory(350–1300ma.s.l.),andalsosouthernmostKurils(KunashirIsland,150ma.s.l.).

Cololejeunea ornataA.Evans,Bryologist41(4):73,1938Description. Plantspalegreenishtobrightandalmost

yellow-green,300–450µmwideand2–5mm long, freely

branched, in loose mats over rocky substrates, bark andotherbryophytes.Rhizoidssparse,colorlesstobrownish,inobliquelyspreadingfascicles,closelyattachingplantstothesubstratum. Stem 40–50 µm in diameter, freely pinnatelybranched, cross section with 5 outer and one inner cellrows. Leaf lobe obliquely inserted and oriented, slightlyconvextocanaliculate,withapexdistinctlycurvedtovent-ralside,obliquelyovatewithacute-apiculateapex,withco-nical mamillose-papillose protrudences over surface, alsoincludingthekeel,keelwithspines12–15µmlong(5–6µmthicknessof terminalpapilla);lobuleconvex,merelyinvo-lute, with conical mamillose-papillose protrudences oversurface, the first tooth 2 cells long, 1 cell wide, conical,withacuteapex, the second tooth reduced,obtuse,1cellhigh.Cellsinlobemiddle10–15×10–13µm,thin-walled,withsmall,concavetrigones, intermediatethickeningsab-sent;oilbodiesoblong,veryfinelypapillose,2–6percell.Stylus robust, 2 cellswide and8–12 cells long, uniseriateend3–5cellslong,withmamillose-papilloseprotrudencesover the margin. Autoicous, freely fertilized. Androecia1(–2)-androus (the second antheridium, if present,muchsmallerthanthefirstone).Perianthloosely4–5–keeled(1–2–ventral,2–lateral,1–dorsal),obovate,ca400×280µm,densely spinoseover surface, exerted for1/2–2/3of thelength, bract lobule almost the same length with lobe.Figures3B,M;10:4,5)

Comment.Thiseasilyidentifiedtaxonisduetoitsspi-nose surfaceof theouter sideof thedorsal lobe andob-liquely oriented (not dorsiventrally appressed) lobe that issometimes somewhat canaliculate. Confusion is only pos-siblewithCololejeunea subkodamaeasdiscussedunderthelatter.

Ecology. Neutro-tobasiphilousmesophyte.InNorthAmerica (south-eastern coast) known from calcareousrocks.InChinareportedonlyfromepiphylloushabitat(Zhu&So2001).Contrary, inJapan it isgrowingas“creepingovermossesandfernsoncalcareousrocks”(Mizutani1961:266).WithintheRussianFarEastthespeciesprefersopento semi-shaded bases of tree trunks (both of deciduousandconiferoustaxa)andneutralandbasic(includingpurelimestone)rocks,includingthosenearstreams,butasideof directimpactof runningwater,andmaybecharacterizedasneutrophilicmesophyte.Itformspurepatchesorgrowingtogether with other Lejeuneaceae (Lejeunea neelgherriana, Lejeunea japonica),Frullania davurica, Metzgeria pubescens, Porella caespitans(Steph.)S.Hatt.,P. vernicosaLindb.,Radula obtusiloba, R. complanata(L.)Dumort.

Distribution.Amongour taxaof thegenus it isoneof themostwidelydistributedspecies.Aside theRussianFarEast,whererelativelycommoninthesouthernflankof thearea,itisquiterareinJapanandKorea(Mizutani1961,Choi 2013). Besides known in North America from thesouthernpartof AtlanticU.S.A. (lowelevations inSouthCarolina,Tennessee,Florida,cf.Theirs2018).Onlytwolo-cations are known inChina (Anhui,Zhejiang, cf.Zhu&So2001).AlsoknownfromPakistan(Furukietal.1993).WithintheRussianFarEast it ischaracterizedbyamoresou thern distribution than C. macounii, although in largescaletheareasareoverlapping.However,thenorthernmostlocality among Cololejeunea is that of C. ornata (49°22'N



inAnujRiver).The latteroccurrencemayhave the relictnaturebecauseitisfromlowelevation(202ma.s.l.)larchforest,notfromhemiborealUssuritaigalikethenorthern-mostoccurrenceof C. macounii.Southwardof KhabarovskTerritory through southern half of Primorsky Territory(likelyshouldbefoundinitsnorthernpart),wherebelow400ma.s.l.,withonelocalityin520ma.s.l.inthemiddle

partof theterritory.BeforeitwasreportedforKhinganskyReserve inAmurskayaProvince (Gambaryan&Cherdan-tseva1998),Komsomol’skyReserve inKhabarovskTerri-tory(Cherdantseva&Gambaryan1989)andSikhote-AlinReserve inPrimorskyTerritory (Gambaryan2001). In alltheselocalitiesitmaybeexpected,however,voucherswerenotfoundinVLA.

Figure 10 Cololejeunea macounii (Spruce)A.Evans:1–shootfragment,androecialbranch;2–lobuleandstylus,ventralview;3–planthabit,dorsalview.Cololejeunea ornataA.Evans:4–planthabit,ventralview;5–lobe,lobuleandstylus,ventralview.1,2–fromP-17–3–12;3–fromPrim-16-26-16;4,5–fromPrim-17-10-16(allinVBGI).Scales:a–500µm,for1;b–100µm,for2;c–1mm,for3;d–400µm,for4;e–100µm,for5


Bakalin

Cololejeunea subkodamae Mizut.,J.HattoriBot.Lab.60:448,1986

Description. Plantsgreenishtodeepgreen,prostratetolooselyascending,well-developed600–700µmwideand3–10mmlong,inloosepatchesoverotherbryophytes,treebarkandrocks.Rhizoidsvirtuallyabsentorsolitary,colorless,obliquely spreading in the area between lobule adnations(twozonesof rhizoidsoriginper leaf pair). Stems freelytosparselybranched,with5outerandoneinnercellrows.Leaf lobesimbricate,whenflattenedintheslideobliquelyovate-triangular, well-developed 350–400× 200–250 µm,erectspreading,slightlyconvex,notorobscurelyturnedtodorsal side,denselypapillose-mamillose throughoutoutersurface,papillaeconicalornearlyso,margindistinctlycre-nulate;ventrallobeobovate,convex(if tolookfromvent-ral side), curved to almost involute inward, keel area andadjacent cell rowswith conical papillae that disappear tothelobuleoutersurfacemiddle,thefirsttoothnarrowlytri-angular,1–2cellswide,2–3cells long(uniseriateend1–2cells long), terminal cell conical, not thickened apically,slimepapillaoninnersurfaceof subapicalcell,thesecondtoothasonlyunclearcrenationornearlysmoothenedcur-vatureor1–celled,withroundedapex.Cellsinlobemiddle12–17µmindiameter,trigonesverysmall,concave,inter-mediatethickeningsabsentorobscure;oilbodies2–6percell,ellipsoidaltoshortlyfusiform,finelygranulate.Stylussmall, 1–2(–3)-celled, easily caducous in older part of shoots.Gemmae discoid, 30–40 µm in diameter. Figures3C,N;9:5–7.

Comment. ThistinyspeciesmaybeeasilyconfusedwithCololejeunea ornata, which shares the conical protrudencesoverthedorsalsurfaceof theleaf lobe.Themainattentionshouldbepaidtothestyluscharacteristicsthatisvestigial(1–2–celled)andeasilycaducousinolderpartof shootinC. sub kodamae,versusmorepersistentandlong(2cellswideandcommonlyover10cells long) inC. ornata.Moreoverthemamillose-papilloseprotrudences aredistributedovertheouter surfaceof the leaf lobule inC. ornata,whereastheir distribution in C. subkodamae is restricted by severalcellrowsadjacentonlytothekeel.PotemkinandAfonina(2008) recorded C. nakajimae for Zabaikalsky Territoryof Eastern Siberia. Despite the latter belonging to subg.Taeniolejeunea(cf.Soderstrometal.2016),itdoesnotshowthecharacteristictraitsof thatsubgenusincluding4–keeledperianthandseriateocelli.Contrary,ocelliareabsentinthespeciesandtheperianth is5–keeled.The latterpermittedMizutani(1961)totreatitasmemberof subg.Cololejeunea.Cololejeunea nakajimae somewhat resembles C. subkodamae fromwhichitdiffersincompletelysmoothoutersurfaceof thelobule(versuswithpapillose-mamilloseprotrudencesinseveral cell rowsnear the keel).Other features, includinglobecellsizeandplantsizeseemtobelargelyoverlapping.Formally thecell size inour specimens from theRussianFar East is rather like the cell size of C. nakajimae than in C. subkodamae.Cololejeunea nakajimae isa rarespeciesof middle Japan (Honshu) thatdoesnot gooutside and thedistribution in the Russian Far East looks less probable(if toputaparttheenigmaticcollectionfromZabaikalskyTerritorymentionedinPotemkin&Afonina2008).

Ecology. Acidophilous mesophyte. In Japan knownfromvarioushabitats,includingtreetrunks,branches,rockysubstrates and living leaves of ferns and vascular plants,whereasinChinathedistributionof thespeciesislimitedbyepiphylloushabitatsinmoist(subtropical?)forestsonly(Zhu&So2001).Ecologyof the species in theRussianFarEast is poorly understooddue to under-collecting. Itoccursoverbasesof Abies nephrolepis (Trautv. exMaxim.)Maxim., where it forms thin pure patches or growingtogether with Cololejeunea macounii. Rarely may be foundovercliffsof neutralreactioninthenorthernmostextremesof thedistribution in theRussianFarEast.Basedontheinformationathanditprobablymaybestbecharacterizedasanacido-toneutrophilicmesophyte.

Distribution.BasicallyaJapanese-Koreantaxonspar-sely distributed throughout Japan (from Honshu to thesouthernmostextremes:IriomoteIsland)andSouthKorea,northwardbecomingveryrareinPrimorskyTerritoryand(probably) SouthKurils aswell as known fromone verydistant locality inCentralAsia (Republicof Tuva), if thelatterreportdoesnotactuallybelongtoanother,probablypreviouslyundescribedspecies.TheepiphylloushabitatsinChinaoccurfrom400to1250ma.s.l.intheprovincesof Fujiang,Guizhou,andZhejiang(Zhu&So2001).Thedist-ributionof thetaxonintheRussianFarEastmaybecalledas‘psudo-enigmatic’.Indeedthespeciesisknownfromonlytwolocalities:BadzhalMts.inKhabarovskTerritory(550ma.s.l.in50°N)andfromLivadijskyRangeof PrimorskyTer-ritory(695ma.s.l.near43°N).Thisdistributionpatternisrathertheconsequenceof under-collectingthantheconse-quenceof realdisjunctions.Thereisoneobscurereportof thespeciesforShikotanIsland(Horikawa1940)thatmaybebasedonamistakeinidentification(cf.Bakalin2007).

P H Y T O G E O G R A P H I C A L R E M A R K SThesouthernflankof theRussianFarEastistheon-

lyplacethatbelongstothenorthernextremeof theEastAsianfloristicprovince–oneof worldwidediversitycen-ters of Lejeuneaceae. Therefore, only remnants of theenormousEastAsiandiversityoccurinourarea.Intotal,theoccurrenceof 11taxaof LejeuneaceaewasconfirmedfortheRussianFarEast:1–Acrolejeunea, 1 – Cheilolejeunea, 1 – Mic ro lejeunea, 4 – Lejeunea and 4 – Cololejeunea.Onlyonespecies,thearctic-alpine,mega-BeringianLejeunea alaskana doesnotoccur in theEastAsianProvince, allother taxaoccursouthwardatleastinJapanand,almostall,inChinaandtheKoreanPeninsula.

Ontheotherhand,thediversityof LejeuneaceaeintheRussianFarEastishighincomparisonwithotherRussianmacro-regions.Twoof eleventaxarecognizedintheareatreatedareknown inRussiaoutsideof RussianFarEast,among themLejeunea alaskanawestwardly reachesTaimyr,Cololejeunea subkodamae–knownfromonelocalityinSouthSiberia (Tuva Republic). Only a few taxa recorded fromotherRussianregionsdonotoccurintheFarEast,inclu-ding Lejeunea cavifolia(EuropeanpartandCaucasus),Lejeunea patens Lindb. (Caucasus), Cololejeunea calcarea (Lib.) Steph.and Cololejeunea rossettiana (C.Massal.) Schiffn. (Caucasus)and Cololejeunea nakajimae S.Hatt.(SouthSiberia).



Themajorityof Lejeuneaceae(exceptof theaforemen-tio ned Lejeunea alaskana)arerestrictedtothesouthernflankof theRussianFarEast.Mostof themareusualcomponentof theflorainthesouthernhalf of PrimorskyTerritoryand,to a lesser extent in the southernmostKurils. The popu-lationsbetween48and54°N(whereLejeuneaceaearecon-spicuousrarity),atleastsomeof them,shouldberegardedashavingarelictnatureduetoobviousdissonanceof thosetaxawithsurroundingvegetationenvironments.Moreover,afewtaxacommonandrelativelyfrequentintemperatefo-rests, including thosewithmany evergreen trees, occur inthenorthernextremityof thearea,andnotatthelowestalti-tudeherebutevenatthehighest!Themoststrikingexampleis Cheilolejeunea obtusifolia occurring in northern-boreallarchforestsat thehighestelevations in theBadzhalMts.,together with e.g.Lejeunea alaskana and Frullania sub arc tica.This phenomenon of unexpectedly northern occurrenceof temperatetaxa isprobablysomewhatrelatedto ‘UmiatSyndrome’describedanddiscussedbySteere(1965,1976).

The Lejeuneaceae occurring in the Russian Far East (aswellasmostof themintheworldwideflora)areverysensitivetoairmoisture(apossibleexceptionmaybeAcrolejeu nea san dvicensis).Thelattercircumstancemayexplaintheoccurrenceof manytaxamostlyoronlyinthemiddleele-vationbeltof PrimorskyandKhabarovskTerritories(thuscontinentmainland)–anareamoistenedbyPacificmon-soon (see Bakalin 2008), contrary to lower elevations insouthernmostKurils. The following examples (basing onspecimensexamined)areillustrative:1)Cheilolejeunea obtusifo lia distributed in (350–)500–1660 in the mainland and15–320ma.s.l.inSakhalinProvince;moreoverthespeciesdoesnotavoidlowerelevationsmuchsouthwardinJapan;2) Cololejeunea macounii – (350–)528–1300 m a.s.l. in themainlandversus150inKurils;3)Lejeunea japonica(35–)300–1600ma.s.l.versus120–300inKurils.

NotabledifferencesareseenbetweenLejeuneaceaeof theinsularregionandof thecontinentalmainland.Forin-stance, theonly insular regionhousesvery sensitive toairhumidityMicrolejeunea punctiformis, a species very sensitiveto air humidity, and the Japanese (in broad sense of thisterm)endemictaxon–Lejeunea otiana.Incontrast,theonlycontinentalmainlandhousesLejeunea neelgherriana (theareaof commonfoggoingfromthesea),probablyCololejeunea subkodamae (if the report for Shikotan is incorrect), andfurthermoreC. ornata, C. japonica and Acrolejeunea sandvicensis.Thetaxarestrictedtothecontinentalmainlandseemstobemore drought tolerant than those limited by insular part.AnotherreasonthatIcanseeisthattheenvironmentsof continentalmainlandinthesouthernRussianFarEastmightbemorestableandmoreheterogeneous,andthusoffermoresuitablemicrohabitatstoaccommodatethemostlytropicalLejeuneaceaewhich,inageneralsense,arealienelementsinthemodernvegetationof theRussianFarEast.

A C K N O W L E D G E M E N T SI am sincerely grateful toMr.MatveiBakalin for line

drawingsforthepresentaccount,Dr.NadyaKonstantinova(KPABG) and one anonymous reviewer for constructivecommentsandDr.JamesShevock(CAS)forEnglishcor-

rectinginthemanuscript.TheworkwaspartiallysupportedRussianFoundationforBasicResearches(17-04-00018).

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Attachment 1SPECIMENS EXAMINED(onlyselectedspecimensarecited;thespecimensoutsideof theRussianFarEastarecitedonlyif theyweredrawnorotherwisementioned).Acrolejeunea sandvicensis:RUSSIA: Amurskaya Province:KhinganskyReserve(49°12'03"N130°34'46"E),343ma.s.l.,leg.CherdantsevaV.Ya.,18.IX.1992(s.n.,VBGI);ZejskyDistrict(54°15'45"N126°50'53"E),340ma.s.l., leg.DudovS.V.,26.VII.2012(Br_12_012,VBGI);Khabarovsk Territory:BadzhalMts.(50°15'43"N134°41'48"E),600ma.s.l.,leg.BakalinV.A., 2.VIII.2016 (Kh-16-22-16, VBGI); Primorsky Territory: Anuchinsky District (43°51'55"N 132°55'33"E), 280 ma.s.l.,leg.BorovichevE.A.&V.ABakalin,9.XI.2014(BB5/11-14,VBGI);AnuchinskyDistrict(43°51'55"N132°55'33"E),280ma.s.l., leg.BakalinV.A.,3.VI.2010(P-3-6-10,VBGI);KavalerovskyDistrict (44°14'58"N135°03'19"E),321ma.s.l.,leg. BakalinV.A., 17.IX.2011 (P-68-21-11,VBGI); (44°14'58"N 135°03'19"E), 321m a.s.l., leg. BakalinV.A., 17.IX.2011(P-68-20-11,VBGI);KhankajskyDistrict (44°50'00"N131°41'45"E),330ma.s.l., leg.BakalinV.A.,2.IV.2011 (P-1-11-11,VBGI);(44°50'52"N131°42'39"E),607ma.s.l.,leg.BakalinV.A.,3.IV.2011(P-2-23-11,VBGI);(42°34'02"N130°48'12"E),150ma.s.l.,leg.BakalinV.A.,29.IV.2016(P-7-9-16,VBGI);KhasanskyDistrict(42°37’15”N131°08’10”E),180ma.s.l.,leg.BakalinV.A.,15.X.2004(s.n.,VBGI);(42°37'10"N131°08'05"E),180ma.s.l.,leg.BakalinV.A.,15.X.2004(140-12-04,VBGI);LazovskyDistrict(43°23'45"N134°11'59"E),356ma.s.l.,leg.BakalinV.A.,20.IX.2012(P-43-15-12,VBGI);OlginskyDistrict(43°44'38"N135°12'24"E),25ma.s.l.,leg.BakalinV.A.,22.IX.2007(P-84-16-07,VBGI);PartizanskyDistrict(43°13'53"N133°24'19"E),600ma.s.l.,leg.GambaryanS.K.,25.IX.1978(s.n.,VBGI).Cheilolejeunea obtusifolia:JAPAN: Miyazaki Prefecture:Nichinan(31°44'00"N131°22'00"E),300ma.s.l.,leg.BakalinV.A.&M.Matsumoto,14.V.2015(J-40-16-14,VBGI).RUSSIA: Khabarovsk Territory:BadzhalMts.(50°18'29"N134°39'52"E),1570ma.s.l.,leg.BakalinV.A.,4.VIII.2016(Kh-17-38a-16,VBGI);BadzhalMts.(50°20'44"N134°39'42"E),1640ma.s.l.,leg.BakalinV.A.,6.VIII.2016 (Kh-19-13-16,VBGI);Primorsky Territory:LazovskyDistrict (43°14'05"N133°44'07"E),350ma.s.l., leg.BakalinV.A.,23.IX.2011 (P-74-38-11,VBGI);LazovskyDistrict (43°14'45"N133°43'02"E),500ma.s.l.,leg.BakalinV.A.,6.X.2006(P-68-12-06,VBGI);LazovskyDistrict(43°14'45"N133°43'05"E),570ma.s.l.,l

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