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( $ Figure  1  )!dxdx { FG'  _XXAndreaeaceae"  38   R(  _j(#XX  38  ڀ"Andreaeaceae(3$ !   | FG'  _XXEncalyptaceae"  98   R(  _i(#XX  98  ڀ"Encalyptaceae(EhCEKQW]cioAutoList21.1.1.1.1.1.1.1. { FG'  _XXThuidiaceae"  210  V1. dYYYY)!dxdx\\\\)!dxdx  d^^^^)!dxdxP Pd````)!dxdx dbbbb)!dxdx d^^)! dxdxeeee)!dxdx&0 d dgggg)!dxdx&P d diiii)!dxdx&p d Pdkkkk)!dxdx&p Pd dmnmn)!dxdx3+0 d d d3+0 d d d } FG'  _XXEntodontaceae"  100   R(  _XXh(#XX  100  ڀ"Entodontaceae } FG'  _XXAulacomniaceae"  40   R(  _>>h(#XX  40  ڀ"Aulacomniaceae ~ FG'  _XXLeptodontaceae"  143   R(  _g(#XX  143  ڀ"LeptodontaceaeGrimmia | FG'  _XXBartramiaceae"  42   R(  _i(#XX  42  ڀ"Bartramiaceae | FG'  _XXFabroniaceae"  101   R(  _i(#XX  101  ڀ"Fabroniaceae R(  _  j(#XX  210  ڀ"Thuidiaceae  FG'  _XXBrachytheciaceae"  46   R(  _f(#XX  46  ڀ"Brachytheciaceae z FG'  _XXTimmiaceae"  211  &c$""0  ~ FG'  _XXFissidentaceae"  102   R(  _\\g(#XX  102  ڀ"Fissidentaceae w FG'  _XXBryaceae"  53   R(  _k(#XX  211  ڀ"Timmiaceae { FG'  _XXFunariaceae"  106   R(  _j(#XX  106  ڀ"Funariaceae } FG'  _XXBryoxiphiaceae"  77   R(  _h(#XX  77  ڀ"Bryoxiphiaceae } FG'  _XXFontinalaceae"  104   R(  _~~h(#XX  104  ڀ"Fontinalaceae z FG'  _XXLeskeaceae"  144   R(  _DDk(#XX  144  ڀ"Leskeaceae | FG'  _XXBuxbaumiaceae"  78   R(  _lli(#XX  78  ڀ"Buxbaumiaceae x FG'  _XXHepatics"  213   { FG'  _XXGrimmiaceae"  109   R(  _j(#XX  109  ڀ"Grimmiaceae { FG'  _XXCampyliaceae"  79   R(  _j(#XX  79  ڀ"Campyliaceae FG'  _XXMosses,IndexandCatalog"  213   z FG'  _XXMeesiaceae"  149   R(  _22k(#XX  149  ڀ"Meesiaceae } FG'  _XXCatoscopiaceae"  82   R(  _nnh(#XX  82  ڀ"Catoscopiaceae | FG'  _XXHedwigiaceae"  134   R(  _i(#XX  134  ڀ"Hedwigiaceae R(  _**\(#XX  213  ڀ"Mosses,IndexandCatalog { FG'  _XXClimaciaceae"  83   R(  _j(#XX  83  ڀ"Climaciaceae } FG'  _XXHylocomiaceae"  135   R(  _66h(#XX  135  ڀ"Hylocomiaceae z FG'  _XXDicranaceae"  84   x FG'  _XXMniaceae"  151   R(  _m(#XX  151  ڀ"Mniaceae y FG'  _XXHypnaceae"  137   R(  _TTl(#XX  137  ڀ"Hypnaceae  k  { FG'  _XXNeckeraceae"  155   R(  _j(#XX  155  ڀ"Neckeraceae  FG'  _XXOrthotrichaceae"  157   FG'  _XXPlagiotheciaceae"  161   R(  _e(#XX  161  ڀ"Plagiotheciaceae ~ FG'  _XXPolytrichaceae"  162   R(  _RRg(#XX  162  ڀ"Polytrichaceae z FG'  _XXPottiaceae"  167   R(  _zzk(#XX  167  ڀ"Pottiaceae FG'  _XXPterigynandraceae"  196   R(  _DDd(#XX  196  ڀ"Pterigynandraceae | FG'  _XXRhytidiaceae"  197   R(  _i(#XX  197  ڀ"Rhytidiaceae } FG'  _XXSeligeriaceae"  198   R(  _h(#XX  198  ڀ"Seligeriaceae { FG'  _XXSphagnaceae"  200   R(  _j(#XX  200  ڀ"Sphagnaceae | FG'  _XXSplachnaceae"  205   R(  _i(#XX  205  ڀ"Splachnaceae ~ FG'  _XXTetraphidaceae"  208   R(  _,,g(#XX  208  ڀ"Tetraphidaceae !  _  o[X` hp x (#%'0*,.8135@8:<H?AXo8xXdd8.J:C:\WAW\CRYPTMS\Mosses2005.wpdJC:\WAW\CRYPTMS\Mosses2005.wpd/    Revised:  March18,2005!     @ 4(AWORKINPROGRES_S_Ԁ)  b XXe    @ 4 GUIDETOTHEBRYOPHYTESOFCOLORADO  0  @{ { &Keysandecologicalnotesbasedonfieldandherbariumstudies   @//<BYWILLIAMA.WEBER 0  @^^2FellowoftheLinneanSocietyofLondon  @R R .ProfessorandCuratorEmeritusoftheHerbarium p @OO2UniversityofColoradoMuseum,Boulder 0 @MMDand  @((<RONALDC.WITTMANN 0 @4Associate,HerbariumCOLO,Boulder   (aWorkingDocument)  0 @$$<March2005version   XxX8xWXdXd8@= TABLEOFCONTENTS    Note:thepaginationchangesaswemakerevisions.Thefamiliesarealphabeticallyarranged. ~ For generathathavebeenmovedtootherfamilies,consulttheIndexandCatalog.Portionsofthe f manuscriptthatareinneedofamplificationareindicatedby***. N Preface(#(#{(#.  j Introduction(#(#{(#.AA  6 HistoricalcollectorsofColoradobryophytes(#(#{(#.OO;    Habitatsofbryophytes(#(#{(#. % ~  GeographyofColoradobryophytes(#(#{(#.ww/ J  Littlerecognizedfactsaboutbryophytedistributions(#(#{(#.D f Somethoughtsaboutecology,evolution,andtaxonomy(#(#{(#.C 2 _Geobotany_Ԁofbryophytes(#(#{(#.S S &  @||>Part1.Mosses z Keytothegenera(#(#{(#.  F Family,genusandspeciestreatments(#(#{(#.  3 b IndexandcatalogofColoradoMosses(#(#{(#.kk3 . Indexbyspecificepithets0 m 0 (# (# m #(#(#{(#.m m ( (# (##  Derivationsofthegenericnames(#(#{(#./ v!  Meaningsofthespecificepithets(#(#{(#.0 B#" SourcesforIllustrations(#(#{(#. ( %^ $ @KK=Part2.Hepatics &*"& Keytothegenera(#(#{(#.  (#( Family,Genus,andspeciestreatments(#(#{(#.gg4 r*%* IndexandCatalogofColoradoHepatics(#(#{(#.5 >,', ЇIndexbyspecificepithets(#(#{(#.c c )  Derivationofthegenericnames(#(#{(#.ee. | Meaningsofthespecificepithets(#(#{(#.0 H SourcesofIllustrations(#(#{(#. '  d @@Appendices  0 AuthorsofBryophytetaxa(#(#{(#. (    Glossaryofbryologicalterms(#(#{(#. , x  Historicalreferences D    1.LeoLesquereux(1884)(#(#{(#./ `   2.TheColoradoCollectionsofA.J.Grout(1914)(#(#{(#.__H ,   3.ThetypecollectionsofT.S.Brandegee(#(#{(#.A    4.ColoradoCollectionsreportedbyT.P.JamesinWheelerExpeditionReport,1878(#(#{(#.G!G!i@(#(#{(# t   5.F.J.Hermann.MossesofRockyMountainNationalPark(#(#{(#.P @   .Additionspaper,1976(#(#{(#.B  \   6.OriginaldescriptionsofmossesdescribedfromColorado(#(#{(#.aaQ (   7.SevilleFlowersMosses:UtahandTheWest(#(#{(#.D    8.Phytogeographicalcomparisons(#(#{(#.7 p!    9.ColoradoBryologicalHotSpots <#"     BoulderMountainPark %X $     MountEvans &$"&     HighCreekFen (#( 0  0 (#(#HallRanch***#(#(#{(#. - (# (## l*%* Acknowledgments(#(#{(#.? ? @(#(#{(# 8,', ЇBibliography(#(#{(#.   (#(#{(#@?Dedicatedto  @L L 6FREDERICKJ.HERMANN,19061987 @  Hisworkwillnotenduntilwehavetracedeachdelicatewebthattiesuswithallthelifeofthe  \ planet.Hisgreatestworrywasthatwewillfinishtheplanetbeforewefinishthatwork.Theworld B andallitsbeautybelongstoallofus,nottopoliticiansandmereideologies.Thatourweaponscould ( eradicatealllife!plants,insects,mammalsasmuchasman!wasaterriblefearforhim,andhe  workedhard,contributinghoursandmoney,againstthatfear.EricHermannin_Vos_Ԁ&_Reznicek_  (1988).    8xXXdWXd8 =xWXdd=@B PREFACE     I(seniorauthor)firstlearnedaboutmosseswhenIwasabouttwelveyearsold.WhenIwas ~ inHighSchoolIbuiltasmallterrariumwithmossesandlichensinit,andentereditinacontest(this d wasbeforethedaysofScienceFairs)attheAmericanMuseumofNaturalHistory.Withthe$25.00 J Iwonasfirstprize,IwasurgedbymybiologyteachertobuyasmallbookbyA.J.Grout,Mosses 0  withaHandLens.Thus,IacquiredmyfirstmosstextbookonChristmasDay,1934,andstillfindit  f usefuleightyyearslater!  L   Mr.GroutwasahighschoolteacheronStatenIsland,andanassociateoftheNewYork    BotanicalGarden.Hespentalifetimebecomingaspecialistonmosses,andeventuallyproduceda    classicthreevolumeworkentitledMossFloraofNorthAmerica,writtenwiththeaidofasmany   specialistsashecouldmuster.MosseswithaHandLenswaswrittenfromthestandpointthatGrout z  feltthat,withproperattentionandguidance,apersoncouldlearntorecognizemostofthecommon `  mossesoftheNewYorkCityRegionwiththeuseofonlyahandlensandhisbook.Thisbookwas F  sopopularthatpeoplebegantobefairlyproficientbutneededmore.Tomeetthedemand,Groutthen ,|  wroteMosseswithHandLensandMicroscope.Thisrequiredmorediligentandcriticalstudy,and b theareacoveredwasgreater.MostprofessionalbiologistsofmygenerationinAmericaowetheir H startstothisremarkableman. .   Grout'slittlemossbooks,nowsadlyoutofprint,wereextremelyusefulintheEasternUnited  States,andhavebeenfollowedbymanyusefulbookscurrentlyavailablethere.Oneofthevery  successfulonesbasedonGrout'sphilosophywasHowtoKnowTheMosses,byHenryS.Conard v 1956),oneofthemostbelovedofteachersofbuddingbryologists.Hewasprofessorat_Grinnell_ \ College,Iowa.Unfortunately,Conard'sbookwasalsodesignedforuseintheeasternU.S.,anddid B nothelpmeatallin1944whenIwascollectingmossesasahobbyintheColumbiaRiverGorge. (x However,thisfactwasnottootragic,becauseitresultedinmycorrespondingwithDr.Conard,who ^ helpedmeidentifymymosses.Perhapsitwasthroughhelpingpeoplelikemethatherealizedthat D hehadtopublishanexpandedversionthatcoveredtheentireUnitedStates.Thelatesteditionisco * authoredbyoneofConard's_protegs_,Paul_Redfearn_,whohasjustnowretired.    OneofGrout'sdiscipleswasSeville( Bill),ortomostofhisstudents, Doc),alongtime   professorofallsortsofbiologyandbotanyattheUniversityofUtah.BillFlowers,overtheyears, r!  emulatedGroutinproducingmimeographedoutlinesofkeystomosses,liverworts,andferns,of X"! Utah.Heamassedalargecollectionaswell,andpainstakinglyproducedmarvelouspenandinkillus >#" trationsforabookonthemossesofUtah.Hedidnotlivetoseethispublished,butafterhisdeath, $$t# HowardCrum,bryologistattheUniversityofMichigan,editeditanditwaspublishedunderthe  %Z $ nameMosses:UtahandtheWest.ForColoradobryologiststhisbookhasexcellentdescriptionsand %@!% illustrationsofnearlyallofourColoradospecies._It___Ԁalsohasoneofthebestintroductionstomoss &&"& morphologyeverwritteninAmerica.AnexcellentaccountofFlowerslifeandworkhasbeen ' #' publishedbyW.H._Behle_Ԁ(1984). (#(   Therearenowseveralexcellentmossflorasavailable,butfewofthemareespeciallypertinent n*%* totheRockyMountainregion.ThePacificNorthwestiswellservedby_Elva_ԀLawton'sMossFlora T+&+ ofthePacificNorthwest;CrumandAndersonhaveatwovolumeMossesofEasternNorthAmerica, :,', andWilliamD.ReesehasrecentlypublishedaMossesoftheGulfSouth.Sharp,Crum,andEckel  havepublishedatwovolumeMossFloraofMexico.Ifanyoneislookingfornewworldstoconquer,  letthemplanaMossFloraofArizonaandNewMexico! |   ThisguidetotheColoradomossesispatternedaftertheexcellentseriesoflittlebookswritten H bythelateProf.HelmutGamsofInnsbruck,Austria,entitled_Kleine_Ԁ_Kryptogamenflora_Ԁvon_Mittel . ~ europa_.Theyare Excursionfloras!fieldguidestomosses,lichens,ferns,fungiandalgaeofCen  d tralEurope,andtheirauthorwasoneofthekeenestfieldmeninEurope.Irememberfollowinghim  J inthefieldandlisteningandlookingashepointedout(sottovocetohimself)allofthevarious  0 specieshesawashepassedby.Thisisonlyoneofthemanywaysonelearnsthelowerplants.      Asacompromiseowingtothedesirabilityofhavinganeasilycarriedfieldmanual,an   ExcursionFloracontainsonlykeysandecologicalnotes.Formoredetaileddescriptionsandillustra x  tionstheypresumetheavailabilityofmorecomprehensiveliterature.Theyalsoexpectacertain ^  amountofgeneralbiologybackgroundonthepartoftheuser. D    Thissecondeditionstilldoesnotaccomplishmyoriginalobjectiveofeventuallyproviding ` aninstructionalintroduction,butavailablebooksandtheplannedMossFloraofNorthAmericafill F thatneed.MuchremainstobedoneinColoradoinrefiningourconceptsofhabitatsanddistribution. , Thefirsteditionspawnedseverallocalamateurswhonowareadeptinrecognizingmosses,and  perhapsmorewillbeencouragedtoenterthefieldwiththisrevision.    Onewhoconstructsdiagnostickeyscanusuallymakeakeythatsatisfieshimself;thatis,he t candistinguishhisspeciesunderthebestofcircumstances,whenallofthenecessarythings(includ Z ingsporophytes)arepresent.Itisquiteanotherthingtoproduceakeythatworksforhisstudents,and @ thathasto workevenifvitalpartsaremissing.Thisisthegreatproblemwithmosses,especially &v inanaridarealikethisone"thatmostofthespeciesarerarelyorneverfoundinfruitingcondition.  \ Thus,thesporophyticinformation,whichoftenmakesidentificationsurerandmucheasier,cannot B beincludedinakey.Thewritingofthekeythusbecomesadifficultart,withinevitablyequivocal ( results.Furthermore,withoutsporophytesitisimpossibletowriteasatisfactorykeytothelarger  groups"familiesandgenera"sothatadifferentmethodhastobetried.Mrs.PatriciaNelson,my  student,workedoutaveryuseful_polyclave_ԀkeyonIBMcardscalledRandomAccessKeytothe   GeneraofColoradoMosses,whichIconsidertobeanindispensableadjuncttotheguide. p!  Unfortunatelytherearenocopiesleft! V"!   AllofthespecieslistedintheGuidearedocumentedbyherbariumspecimensinthe "$r# herbarium(COLO)oftheUniversityofColoradoMuseum,whichnowcontainsover112,000 %X $ specimensofbryophytes.Iwelcomecommentsandcriticism,andamalwaysinterestedinaddingto %>!% thecollectionsthroughcontributionsfromcollectors. &$"&   Althoughthisdistillationrepresentsmorethanfiftyyearsofsporadicresearch,theworkis (#( nowonlywellbeguninearnest.Thestonesarelaid,butthebuildinghasnotassumeditsfinalshape. )$) Wethinkweknowwhatgrowshere,buttherewillbeadditionsassoonasactivefieldworkbymany l*%* studentsbegins.Wehavesomenotionsofhabitats,buttheyareprimitiveandneedrefinement.Every R+&+ observercomestothefieldwithadifferentpairofeyesandadifferentcoreofexperience,andthere 8,', havebeentoofewofusuptonow.Ishallbesatisfiedifthisguidewillservetobringbryologywithin  thecompetenceofstudentsandecologicalresearchers,andserveasaguidetovisitingbryologists  fromabroad,untilsuchtimeasamoresophisticatedworkismadepossible. |   Thismanuscriptisaguidebookinprogress.Wearemakingitavailableonthiswebsiteto H makethesectionspartsavailableastheynearfinalform.Manyofthesmallerfamiliesarereadyfor . ~ use,butsomeofthelargerormoredifficultonesarebeingworkedoncontinuously.Itishopedthat  d thisversionmightbeusefultoamateurs,students,_amd_Ԁprofessionalecologists.Thismanuscriptis  J anidentificationguidewithmoredetaileddescriptionsthanareusuallygiveninkeys,supplemented  0 bynotesonfieldaspect,ecologicalniche,andotherinterestingobservations.Acollaborative    productioncontributedbyahostofprofessionalbryologistsisinpreparation,whichwillprovide    detaileddescriptionswithillustrationsofallthespecies.Tentativetreatmentsareavailableonthe   NewYorkBotanicalGardenwebsite<4 O  5  _www.nybg.org/bsci/bfna>.66DJDsma  7 !xD x    D  @= SCOPEOFTHEBOOK     Thisisnotanintroductiontothestudyofmossesandliverworts.Infact,itisnotaprimerfor ~ beginners,butasourcebookandguideforpersonsalreadyliterateinthesubject,especiallythose d whocometoColoradofrompreviousexperiencewithbryophytesfromotherpartsofNorthAmerica J andabroad.Elementarybooksonthelifehistories,morphology,andecologyofbryophytesare 0  availableelsewhere.Thisbookconcentratesonthetaxonomyandecology,andthehistoryand  f evidencefortheoccurrenceoftheentiremossandhepaticfloraofColorado,fromthetimeofthe  L historicalexpeditionsofthenineteenthcenturytothepresent.  2   Ourintentionhasbeentoseeifwecanbringtheknowledgeofoccurrenceofthespeciesup    tothepointofdiminishingreturns.Weareconvincedthatthereismoretodiscover,andthatour   understandingofgeographicaldistributionandofhabitatsisstillveryimperfect.Weareespecially z  interestedindrawingattentiontotheprecisemicrohabitatsofthespecies,forfieldworkinbryology `  isonlysuccessfulwhenthemindisprepared.Wecanbegratefultowhatevercreativeforcethereis F  onourplanetthatbryophytes,nolessthanotherplants,arenotdistributedatrandom!Iftheywere, ,|  ourtaskwouldbehopeless. b   Wearealsoawarethattherearewaysbywhichwecantrainoureyestoseedistinctive . charactersofbryophytesthatenableustorecognizethedifferenttaxainthefield.Thesearewhatthe  ornithologistscall flashcharacters.Suchcharactersarenotalwayseasytodescribenorarethey  infallible,butasonegainsexperience,theybecomeindispensableforfieldidentificationandcan  preventexcessiveaccumulationofcollectionsofspeciesthatwearenotparticularlyinterestedin v crammingintoourlimitedspaceinhomeorlaboratory.Beingsuccessfulinfindingwaystorecognize \ Ceratodonpurpureus,thatmostpervasiveherbariumweed,becomesavaluableattributeinthefield B botanist. (x   Learningtorecognizehabitatisveryimportantindevelopinganunderstandingoftherelative D fidelityofbryophytespeciestoacidic,calcareous,mineralized,acidicorbasicsubstrates,.Westill * knowverylittleaboutthisandwonderwhetherourlocalsituationagreeswiththatofthesamespecies  inScandinaviaorAustralia.    Coloradodoesnotappearatfirstglancetobeaveryinterestingplaceforbryophytes"much r!  toodryandsteep.Onedoesntseemanymossesbesidethesuperhighways,andthenonlyafewvery X"! conspicuousandcommononesonseepingroadcuts!Also,thepacetakenbyhikersdoesnotlend >#" itselftooursubject.Whileavascularplantcollectormaycovertwentymilesinaday,thelowly $$t# bryologistmaynotstraymorethanafewhundredyardsalongthetrailbeforebeingforcedtostop  %Z $ foranhourorsoonhandsandkneesbesideaninitiallyunlikelylookingsite.Thus,ifweweretoplot %@!% onamapthelocationsofcollectionsofbryophytesinColorado,thedotswouldfollowtheroads. &&"&   Wemakenoapologyforthetechnicaldifficultyofidentificationofbryophytes.Anyonewho (#( haahandlenscan,withsomeguidanceinthefield,learnmanyofthecommonandconspicuous )$) species,Forthispurposewehaveproducedaprimerforthoseforestersandwetlandmanagerswho n*%* needtobeabletodothistoalimiteddegree..Foramateurs,beingabletorecognizesomecommon T+&+ bryophytesaddstotheamateursjoyinthefield.Infact,Groutsoldbook,MosseswithaHandLens, :,', demonstratesthatitcanbedone.    However,inalltaxonomicstudiesbutparticularlyinthesmallerlifeformsmoresophisticated | equipmentisneededforidentificationrequiringhighmagnificationanddissectionorsectioningof b leavesandstems.Also,nowthatgreateremphasisisbeingmadeonsubtlecharacteristicsand H biochemicalinformation,itisbecomingdifficultforamateursandothernonprofessionalbryologists . ~ todothiswork.Uptoapoint,itismostlyexhilaratingfun,butsuddenlyonehitsastonewall.This  d happenseventoprofessionalswholacknotonlythemoneyforphysicalequipmentbutalsothe  J _indispensible_Ԁreferencebooks,especiallyastheolderonesarebecomingunavailableorarebeing  0 reprinted.These,aswellasthenewbooks,areonlyavailableatastronomicprices.      Thebasicequipmentforaseriousamateurincludestwomicroscopes,onecompoundforhigh   magnificationsandpreparedslides,andonestereoscopicformagnificationsupto100xforgross x  examinations.Dissectingneedles,razorblades(doubleedged bluebladesfromwhichsliverscan ^  bebrokenofftoserveasknives),slidesandcoverglasses,apropaneburnerforheatingdry D  specimensinwater,andafewchemicalreagents,areallnecessary.Dependingonthestudents *z  affluence,afiberopticlightsourceandadigitalcameramountedonthecompoundscopewillmake ` lifeeasier.  F @? INTRODUCTION     Coloradobryophyteshaveneverbeentreatedfully.Theearliestpaperattemptingtolistthem ~ isthatofLeoLesquereux(1874).A.J.GroutvisitedColoradoonceandpublishedashortpaper d (Grout1916)onhiscollectionsnearTolland,inGilpinCounty.FredJ.Hermann(1970,1987) J publishedafewpapers,oneonRockyMountainNationalparkandtheotheronadditionstotheflora. 0  HermannslifeandcareerisreviewedbyVossand_Reznicek_Ԁ(1988).CraftandCraft(1952)taught  f attheAdamsStateCollegeat_Alamosa_Ԁandpublishedathreepagepaperlistingbycounty40mosses  L theyhadcollectedinsouthernColorado,butcitednospecimens.Theyimpliedthatthecollections  2 wereinthatinstitution.H.S.Conard,Frederick_McAllister_,andothershavecollectedmosses    _sporadially_Ԁbutnopublicationsarebasedontheircollections.      Therealfounderofthebryophyteherbarium(COLO)attheUniversityofColoradohastobe z  Geneva_Sayre_Ԁ(_Pfister_Ԁ1993).ShecamefromMenlo,Iowa,andwentto_Grinnell_ԀCollege,whereshe `  cameunderthewingofHenryS.Conard.Whenshegraduated,duringtheGreatDepression,she F  couldnotaffordtogotograduateschool,soDr.Conardinvitedhertostayanotheryearat_Grinnell_ ,|  sothathecouldcontinuetobehermentor.In1934Conardarrangedforhertospendthesummerwith b Dr.Groutin_Newfane_,Vermont.WithGroutsheworkedonthepreparationofthetextforthe H Splachnaceae,Timmiaceae,andAulacomniaceaefortheMossFloraofNorthAmerica.In1935she . receivedtheMastersDegreeattheUniversityofWyoming,herthesisbeingtheworkjustmentioned.  In1935shewentontogetadoctorateattheUniversityofColoradounderJosephEwan.    JimmyhadnomentorsintheregionsoshekeptintouchwithConardandGrout.The v difficultiesofwritingamossfloraforColorado(herthesissubject)wereimmense.Therewasno \ mossherbariumhere;thelibrarywaspoor.Groutsflorawasonlytwothirdspublished.Nevertheless, B shedidwriteathesis.Shewasneververyproudofit,throughnofaultofhers;theinformationwas (x justnotreadyforher,andfieldworkmusthavebeenextremelydifficultwithoutacar.Duringthis ^ timeshestudiedthevascularplants.ItissaidthatshewroteaspringfloraoftheLaramieareaand D amountainfloraofBoulderCounty;thesewereprobablyneverreallypublishedandevidentlyno * longerexist.Butshedidleaveasmallcollectionofherbryophytes,some2,000specimens,which  IdiscoveredinacardboardboxsoonafterIarrivedoncampus.Thiswasourbeginning.    Geneva_Sayre_ԀheldaninstructorshipintheBiologyDepartmentandtaughtattheUniversity r!  Campfrom19381941beforetakinganAssistantProfessorshipatRussellSageCollegeinTroy,New X"! York.ThereshehadtoteachalmosteverythingintheBiologyDepartment(exceptbryophytes!)and >#" spentherentirecareerthere,becomingheadofthedepartmentin1946.Shetookearlyretirementin $$t# 1972andtookaparttimepositionatHarvardinthe_Farlow_ԀLibraryandCryptogamicHerbarium.  %Z $   Becauseofthepressuresofhervariousacademicdutiesandthelackofanherbarium,shewas &&"& neverabletopursueacareerinthetaxonomyofbryophytes.Insteadsheembarkedonseveral ' #' bibliographicprojectsthathaveproducedextraordinarilyimportantbasictoolsforresearchin (#( botanicalbibliography(see_Sayre_Ԁ19571975). )$)   IgottoknowGeneva_Sayre_Ԁatnationalmeetingsandalwayslookedforwardtofindingher T+&+ there.Shewasabrightyoungladyandfullofideasandconversationtopicsinbryology.Inher :,', biographythereisnothingsaidaboutherhavinganythingtodowiththeformationofthebryological  herbariumatBoulder,butIfeelweoweheragreatdealforwhatsheachieved,anditisagreattribute  tohermentorsthatshefulfilledhermissioninlifesoabundantly. |   FredHermann(seeVoss&_Reznicek_Ԁ1988)cametoFortCollinsin1970,withtheherbarium H oftheU.S.ForestService,wherehehadservedinWashington,D.C.andbecametheleading . ~ AmericanspecialistonthegenusCarex.Fromthattimeonuntilhisdeathin1987hedevotedhis  d sparetimetocollectingbryophytes,mostlyinRockyMountainNationalPark,recruitinganow  J famousprofessorofphilosophy,HolmesRolstonIII,totheenterprise.FredandIfrequentlywentinto  0 thefieldtogetherandwehaveatherbariumCOLOduplicatesofmostofhisColoradocollections.    HisprincipalcollectionwenttotheUniversityofMichigan,wherehestartedhisbotanicalcareeras    afreshmanstudentinthe1920s.Itwasagreatprivilegetohavehadhiscompanionshipduringthose   years. x    FredwasfortunatetofindacompatiblecolleagueattheStateUniversityatFortCollins" D  HolmesRolstonIII,aphilosophyprofessor.Wearefortunatethatthesetwomenspentmanydays *z  collectingbryophytesinandnearRockyMountainNationalPark.Holmeswasjustthecompanion ` Fredneeded,foritisverydifficulttobeabryologistandhikealoneinsafetyinthewildernessduring F onesdecliningyears.Holmesprovedtohavenotonlystamina,butanexcellenteye,andcollecteda , numberofexcitingspecimens.However,whenFreddied,Holmeswasabletodevoteallofhistime  tohisfavoritestudy"theapplicationofreligiousteachingsandethicstoenvironmentalconservation.  Hepublishedmanysignificantpapersandtraveledworldwideonthe lecturecircuit,andin2003,  heearnedthe_Templeton_ԀPrizeforProgresstowardResearchaboutSpiritualRealities,joiningMother t TheresaandAlexanderSolzhenitsyninthelistofnotablerecipients.Webryologistscanbevery Z proudthatsuchanindividualhasworkedinourmidst. @   HepaticsaremuchlesswellknowninColorado.AlexanderEvanspublishedalistofColorado  \ Hepatics(Evans1915),butnevervisitedColoradotoseetheminthefield.T.C.Frye&LoisClark B publishedaworkontheNorthAmericanHepatics(Frye&Clark1937,1946)andcitedspecimens ( fromColorado,butlikewisedidnovisitthestate.Theirworkatleastpermittedustocompileacheck  listfromtheliterature.WhileRudolphSchusterpublishedagiantsixvolumeworkontheHepatics  ofpartofEasternNorthAmerica(Schuster19661992),hedidnotincludeobservationsonthe   westernAmericanflora.AtthepresenttimeWon_Shic_ԀHong,CollegeofGreatFalls,ispublishing p!  papersonthewesternAmericanleafyliverwortsandhasexaminedourcollectionsandprovided V"! identifications(Hong19862000)whichhaveprovedveryhelpfultous,buttheAmericanWestneeds <#" moreactiveworkersinthefield. "$r#   ThereisnosingleguidethatcoversalloftheColoradomosses.SevilleFlowers,Mosses:Utah %>!% andtheWestlacksseveralofourgeneraandspecies,asdoesCrum,Steere,andAnderson,Mosses &$"& ofEasternNorthAmerica;Sharp,Crum,andEckel,TheMossesofMexico,treatsseveralColorado ' #' speciesthatotherwiseareunknowninNorthAmerica.SteeresMossesofArcticAlaskahasmostof (#( thedisjunctarcticspeciesbutnokeysareprovided.Themostusefulbookcoveringourfloraactually )$) isNyholmsMossFloraof_Fennoscandia_,whichhasbeenmostusefultousbecausesomanyspecies l*%* arecommontonorthernEuropeandtheRockyMountains.Andlastly,afewofourmosseshavebeen R+&+ addedtoourfloraastheresultofrecentmonographssuchasBlomsrevisionoftheSchistidium 8,', apocarpumgroupinNorwayandSweden,whichhasaddedovertenmorespeciesinthisgroupthan  wasknown.AfewmossesarewidelydisjunctfromMiddleAsia,oneofthem,Didymodon  anserinocapitatus,havingbeendescribedlittlemorethanadecadeago.Thus,thepresentmanuscript | shouldbeofgreatvalueinprovidinginformationaboutspeciesmissingfromtheotheravailable b books. H   ThefirsteditionofthisGuidetotheMossesofColoradowaspublishedinJuly,1973,as  d OccasionalPaperNo.6oftheInstituteofArcticandAlpineResearch.Inthe28yearsfollowingthis  J publication,numerousadditionsandchangeshavebeennecessitated.  0 _      @/HISTORICALCOLLECTORSOFCOLORADOBRYOPHYTES     TherewerenobryologistsinColoradoduringthehistoricexploratoryperiodofthenineteenth ~ century(theUnitedStatesGeologicalSurveys(seeRothrock,1878)andtheHaydenSurveys(see d Foster,1994).NotthattheLewisandClarkexpeditionwouldhavecollectedmanymosseshadit J marchedthroughwhatisnowColorado.Onlytwospecimensofbryophytes,Hypnum(=_Kindbergia_) 0  oreganaand_Bazzania_Ԁtrilobatawereknowntobecollected(Moulton1999),bothofthemare  f exceedinglycommonnorthwesternspeciesthatmerelycouldhavebeenseparatedfromsomevascular  L plantsrootsduringthepreparationofspecimens.Therewerehardlyanybotanistseither;mostwere  2 medicalmenattachedtoexpeditions.Itisnosecretthatbotanistsarethelastinlineforpositionson    generalexploringexpeditions.Theyareusuallyheldhostagebythedemandsofthemoreimportant    members.Theircollectingspotswereusuallyaccidental,suchasreststopsalongtheway.Whenthe   geographers,surveyors,military,geologists,andzoologistswantedtomovetheircamp,thebotanists z  hadtogoalongwiththem.Eventhosewhohavebeenidentifiedas botanistscollectedonlyvascular `  plants.Thelistissmall;mostweremale. F    Fortunately,inthehistoricperiodthereweretwogreatbryologistsinAmerica,whomay b rightlybecalledjointly,the fathersofAmericanbryology"WilliamStarlingSullivantandLeo H Lesquereux(seeRodgers1940).Thankstotheeffortsofthesemenandtheircollaborators,Thomas . P.James,and_Coe_ԀFinchAustin,theyputtogetherthefirst comprehensivemossfloraofNorth  America(Lesquereux&James1884)).  0  e _Biddlecome_,MissHannahJ.AmateurbryologistfromSpringfieldandColumbus,Ohio.Bryum v _biddlecomei_,fromAlma,ParkCo.Andrews,inGrout(MFNA2:224.1935),wrote: The \ originalcollectorisnotknownwithcertainty.Miss_Biddlecome_Ԁ[whoneverwasinColorado] B hadreceiveditfromMrs.Haines,whomayhavereceiveditfromsomeoneelse.(x(#(# 0  e Brandegee,_Townshend_Ԁ_Stith_Ԁ(18431925).CivilengineerwiththeAtchison,TopekaandSantaFe D Railroad,assignedin1871toCanyonCityarea,becamethecountysurveyorofFremont * County,andissaidtohave laidoutthetownofFlorence.Heservedasbotanistonthe  HaydenSurveyin1875andprovidedanearlyandveryusefuldescriptionofthefloraof  southwesternColorado.Hisbryophytescamefrom S.W.Coloradoor within100miles   ofCaonCity.Hewastheonlyresidentbotanistthatcollectedanymosseshere.Afewof r!  thefragmentaryspecimensonwhichnewspecieswerebasedhavenotbeenlocatedagain. X"! BrandegeelaterwentontohaveadistinguishedcareerinCalifornia,wherehewasabotanist >#" attheUniversityofCalifornia,Berkeley.$$t#(#(# 0  e Conard,HenryShoemaker(18741971).ImportantAmericanbryologistandteacherofbryologists, %@!% professorat_Grinnell_ԀCollege,collectedsomemossesnearBuenaVistainthesummerof &&"& 1941.HisherbariumisatIowaStateUniversity.' #'(#(# 0  e Cooper,David.Contemporaryplantecologist,specializingintheflorasofColoradohighaltitudefens )$) andresponsibleforimportantcollectionsofSphagnumandAmblystegiaceae.n*%*(#(# 0  e Craft,JamesH.professoratAdamsStateCollege,Alamosa,publishedalist(1952)ofmosses :,', collectedinseveralsouthernColoradocounties,mostlydeterminedbyConard.These  evidentlyhavenotbeenpreserved.(#(# 0  e Crandall,C.S.(18521929).ProfessorofBotanyatColoradoAgriculturalCollege,collectedsome b mossesinwhatisnowtheRooseveltNationalForestin1894.H(#(# 0  e _Downie_,TimothyCampbell,18301875.Nicknamed Major_Downie_,hecollectedsomemossesin  d Coloradoin1868,atTwinLakes,withtheHaydenSurvey(Ewan1981).Accordingto  J Hayden(1869),theexpeditioncameintoColoradofirstfromCheyenne,visitingpointsinthe  0 outerFrontRangetoColoradoSprigsandRatonPass.ItreturnedtoColoradofromSanta_F_,    uptheColoradoRiver,throughtheSanLuisValley,over_Poncha_ԀPass,throughSouthPark,    andintotheupperArkansasRiverValley.PresumablyTwinLakeswastheirlastcampbefore   returningtoDenver.ThecollectionsweretobeturnedovertotheSmithsonianInstitution,but x  wehavebeenunabletofindthem.^ (#(# 0  e Flock,JoAnnW.(***)StudentatUniv.ofColorado,collectedandpublishedontheinventoryand *z  ecologyofalpinelichensandbryophytesonNiwotRidge,BoulderCounty(Flock1978).Also ` madeinventoriesofthemossesandlichensofCapePrinceofWales,Alaska.F(#(# 0  e _Goodding_,L.N.(18801967),Wyomingbotanist,studentof_Aven_ԀNelson,collectedafewmosses  inRoutCountyintheParkRange(Ptychostomumpallescens,No.1781[COLO])inAugust,  1903._Goodding_smaincollectionswereattheRockyMountainHerbarium,whose  bryophytesweregiventotheUniversityofTennessee,wheretheywerelaterdestroyedina t firethatdestroyedtheherbarium.Z(#(# 0  e  (#(# 0  e Grout,AbelJoel,(18671947).GroutmadeashortvisittoColoradoin1916,andcollectedinthe &v vicinityoftheUniversityofColoradossummercampatTolland,GilpinCounty.See  \ appendix,andSteere(1948).B(#(# 0  e Haines,Mrs.MaryParry(18261884).Anaturalistofeclectictastes,custodianofthePaleontology  Dept.ofamuseuminRichmond,Indiana,andanamateurhorticulturist.Althoughshenever  visitedColoradoshewasresponsibleforfunnelingseveraloddspecimens[cf.Orthotrichum   _hainesiae_)collectedbyherfriends,Mrs.E.J.SpenceandT.S.Brandegee,tobryological p!  specialists(Flowers1942).V"!(#(# 0  e Hall,Elihu(18201882).AfarmerandamateurbotanistinAthens,Illinois.CollectedwithParryin "$r# Coloradoin1862.AlsocollectedplantsinTexasandOregon.%X $(#(# 0  e Hermann.FrederickJ.(19061987),receivedhisdoctorateattheUniversityofMichigan.Hiscareer &$"& wasspentattheUnitedStatesForestServiceHerbarium,wherehisspecializationwasthe ' #' genusCarexandVicia.HeretiredtoFortCollins,Colorado,wherehecollectedafineseries (#( ofbryophytes,especiallyfromtheFrontRangeandRockyMountainNationalPark.His )$) careercollectionsareattheUniversityofMichigan,withmanyduplicatesattheUniversity l*%* ofColorado.SeeVoss&Reznicek(1988).R+&+(#(#  8,', 0  e Holmen,Kjeld(19211974),DanishbryologistwhodiscoveredOreasmartianaonMountEvans.  SpecialistinbryophytesofGreenland.See_Warncke_Ԁ_(1975)._(#(# 0  e Holzinger,J.M.(18531929),GermanbornMinnesotan,collectedbryophytesinColoradoin b ArapahoandPikeNationalForestsin1896,andhisColoradocollectionscontributedtohis H publishedexsiccati,Musci_Acrocarpi_Ԁ_Boreali_Ԅ_Americani_.. ~(#(# Holzinger,Marie.CollectedinBoulderCountyin1892.  J 0  e 0(#(# e Jamieson,David,contemporarybryologist,professoratFortLewisCollege,Durango,andspecialist    inthemossfloraofthesouthwesternSanJuanMountains.AstudentofWilfredSchofield,    Univ.ofBritishColumbia,andmonographeroftheAmericanspeciesofHygrohypnum. (#(# 0  e 0(#(# e Kiener,WalterB.(18941959),SwissbornColoradomountaineer,collectedonLongsPeak(1938), ^  wherehewasaclimberandguide.Hisbryophyteandlichencollectionswereacquiredbythe D  UniversityofNebraskain1960.KienerwasinvolvedinawinterclimbofLongsPeakon *z  Jan.1012,1925,inwhichhiscompanion,Agnes_Vaille_,diedinafall.`(#(# 0  e Lehr,Paula,contemporaryamateurbotanist,collectedbryophytesintheGunnisonBasinandElk , Mountains,andstimulatedinterestinmossesamongamateursandForestServicestaffinthe  localarea.(#(# 0  e Nelson,LawrenceT.(18621932).Primarilyamycologist,collectedforthe_U.S.D.A._Ԁasaforest t agentoccupiedinlocatingpoisonousplantsinColoradoandelsewhere.Hecollectedafew Z mossesfromtheGunnisonarea.Nelsontaughtatanextraordinarynumberofcollegesand @ universitiesinmanystates.Hisbryophytecollections(19021910)areatDukeUniversity. &v SeeGier(1953). \(#(# 0  e Nelson,Patricia(1940),contemporaryamateurbryologist,LibrarianattheUniversityofColorado ( MedicalSchoolinDenver,studiedthemossesofClearCreekCanyon,andforhermasters  thesispreparedaveryusefulRandomAccessKeytotheGeneraofColoradoMosses.An  amateurbryologist,sheiscreditedwiththediscoveryoftheonlycollectionofLeptodon   smithiiinNorthAmerica.p! (#(# 0  e Parry,CharlesChristopher(18231890). Duringthatyear[1865]RockyMountainmosseswere <#" receivedfromDr.Parry,whoafewyearsbeforehadnamedColoradosRockyMountain "$r# peaksforDr.GrayandDr.Torrey)(Rodgers1940).OnlyBartramiastrictawasspecifically %X $ attributedtoParrybyLesquereux&James(1884).SeeWeber(1997).%>!%(#(# 0  e Porter,ThomasConrad(18221901).Botanistandclergyman,wasattachedtotheHaydenSurveys ' #' from18691874((Foster1994).Hismosscollectionswerefewandincidentaltovascular (#( plants.ProfessoratLafayetteCollege,Lancaster,Pa.Whatcollectionsofhisremainareatthe )$) PhiladelphiaAcademyofNaturalScience.l*%*(#(# 0  e Rolston,Holmes,III.(***).Contemporaryamateurbryologist,DistinguishedProfessorof 8,', PhilosophyatColoradoStateUniversity, fatheroftheconceptofenvironmentalethicsas  amodernacademicdiscipline.CollectedbryophyteswithFredHermannforseveralyears,and  isanexcellentfieldbryologist.|(#(# 0  e Rothrock,JosiahTrimble(18391922).AstudentofAsaGray,andabotanistandsurgeononthe H WheelerExpedition.HewasthesupervisorofJohnWolf.Manyvascularplantsbearhis . ~ name;whetherhereallycollectedbryophytesisquestionable. d(#(# 0  e _Sayre_,Geneva(19111992),CollectedmossesintheFrontRangeandinMesaVerdeNationalPark  0 andwrotebutdidnotpublishaMossFloraofColorado.Didextensivebibliographicwork    oncryptogamicexsiccati.See_Pfister_Ԁ(1993).  (#(# 0  e Spence,Mrs.E.Jane.ReportedashavingcollectedatManitouandvicinityinJuly,1897(Ewan x  1981).PossiblythecollectorofNeckeradouglasii?^ (#(# 0  e _Stiverson_,ClareL.Bryologystudent,producedaMastersThesisfortheUniversityofDenverin *z  1951:ApreliminarysurveyofthemossesintheMountEvansarea,45pages.Someofher ` collectionsareinherbariumCOLO.accessionnumbersB92639284.F(#(# 0  e _Wikel_,PatricianeeThomas(***).CollectedhepaticsintheFrontRangeandSanJuanMountainsin  the1970s.CollectionsareatCOLO.(#(# 0  e Wolf,John(18201877).Memberofthe1873Wheelerexpedition.AssistanttoRothrock.Several t vascularplants(cf.Ribeswolfii)bearhisname.Itisquestionablewhetherhehadanyreal Z interestinbryophytes.@(#(#   In19***theUniversityofColoradohostedaseriesoffieldtripsduringthefirst_ICSEB_  \ meetinginBoulder,Alargenumberofbryologistsattended,includingWilliamC.Steere,Lewis B Anderson, (    @: Habitatsofbryophytes   @8Wetmeadows,BogsandFens. ~   Wetmeadowsontheeasternplainstheseverypoorinmosses,butusuallysupport J Drepanocladusaduncus. 0    Carrsarewillowdominatedwetlandscommononmountainfloodplains.Thesearenotvery  L productiveofbryophytesbecauseoferosivescouringbywaster,anddensityofCarexstands.  2 However,atthebasesofwillowshrubs,acharacteristicclusterofmossspeciesincludesClimacium    dendroides,Helodiumblandowii,andTomenthypnumnitens.      Coloradohasnotruebogs.WhathavebeencalledbogsinColoradoarereallyfens._McQueen_ z  (1990)describedthedifferences.Weparaphrasesomeofthesestatementstoapplytoour `  mountainousregion.Bogsarewetlandsthatareverypoorinnutrientsbecausemostoftheminerals F  areobtainedfromprecipitationratherthanfromgroundwater.Theyarealsoreferredtoas ,|  ombrotrophicpeatlands,meaningsimplythatallnutrientscomestrictlyfromprecipitation.Theyare b usuallycharacterizedbyapHof4orless.Sphagnumisthedominantvegetationofabog. H   Fensareverysimilartobogs,buttheyreceivetheirnutrientsfromthesurroundingground  waterandarewetter.Manyofthesameplantsthatarefoundinbogsarealsofoundinfens.Fensare  oftenreferredtoasminerotrophicpeatlandsbecauseifthegreaterinfluxofnutrientsfromground  waterThetermfenmaybesomewhatmisleading,becausethecategoryincludessomediverse v habitats.Forexample,thewordfenisusedtodescribesomealkalinewetlandsthataredominatedby \ sedgesandgrassesandveryfewpeatmosses. B   Poorfensgenerallyobtainmorewaterfromprecipitationthanfromrunofffromelevated ^ places,andtheirpHishigher(5to6.5).HighCreekFen,inSouthPark,maybesocharacterized. D Richfensoccurinlevelareasintowhichwaterdrainsfromhigheraltitudesandflowsthroughand * outalthoughataratherslowrate.ChattanoogaFen,intheSanJuanMountains,isanexample.    Theextensivewillow/sedgemeadowsalongthesubalpinestreamsdonotholdwaterforlong   periodsandmaybecomequitedryseasonally.Therelatively_meagre_Ԁnumberofbryophytesare r!  commonlyrestrictedtotheslightlyraisedbasesofwillowclumps. X"!   Sitesthataredominatedbyrunningwaterarethemostvariedhabitatsinthemountains,and $$t# arenoteasilycharacterized.Theyvaryfromslowlyflowingorintermittentstreamletstorushing  %Z $ cataracts.Someverycommonspecies,suchasBrachytheciumrivulareHypnumcupressiformerun %@!% thegamut,whilecertainspeciesrequiresomesubtlecharacteristicsofthestreamthatarenoteasily &&"& pinpointed. ' #' @<Cliffsandboulders )$) *** T+&+  :,', @]]8Walls,lawns,andsidewalks    AgreatdealhasbeenwrittenaboutthemossvegetationofwallsinEurope,muchlessin | America.InColorado,atleast,ourbuildingsarenotveryold,.andwelackthestonewallsseparating b agriculturalfields.Ourdryclimatedoesnotencourageverymuchwallvegetation. H   Nevertheless,wallmossescanbefoundhere,especiallyonthebuildingsoftheUniversityof  d Colorado,wherenativestone,particularlytheLyonssandstone,hasbeenusedforfacings.Mosses  J occurparticularlyonmortarinthecracksbetweentheflagstonelayersoneastfacingwalls.There  0 arealsoretainingwallsonthecampusthatsupportmosses.Whentheseoccurinareasthatareshaded    thecumberofspeciesincreases.      Someofthespeciesthatarecommononmortarinhorizontalcracksare:Bryumargenteum, x  Barbulaconvoluta,Ceratodonpurpureus,Coscinodoncalyptratus,Didymodonrigidulus,Grimmia ^  anodon,G.pulvinata,Orthotrichumhallii,Ptychostomumangustifolium,Schistidiumconfertum, D  Syntrichiavirescens,andTortulamuralis.Shadedretainingwalls,inadditiontosupportingthese *z  species,havesomespeciesofmoremesicpreferences,suchasBrachytheciumerythrorrhizon, ` Rosulabryumflaccidum,Tortulaatrovirens,andT.mucronifolia.. F   LawnsthatareoverirrigatedorpoorlydrainedsupportmostlyBrachytheciumerythrorrhizon  andCeratodonpurpureus.Sidewalkcracks,astheybecomeeroded,arecommonlyfilledwithBryum  argenteum.FormanyyearstheUniversityofColoradohadasystemofshallowconcreteditches  collectedtoamajorirrigationcanal.Theseditcheswereusedtofloodthecampusmallandarenow t nolongerusedanddry,butreceiveenoughseepagemoisturethattheirverticalsidesareoften Z plasteredwithmatsofAmblystegiumvarium!Adrainpipeonthecornerof_Denison_ԀBuildingwatered @ thesurfaceatitsbaseenoughtosupportanicecolonyofGemmabryumsubapiculatumuntilthe &v FacilitiesManagementpeopledecidedtocleansethebuildingofasbestosandbuiltatemporaryshack  \ overthesite.Mostofthemossdisappearedbutisslowlycomingbackdespitethelackofwater! B @>Otherhabitats    Alkalineflatsareseasonallywetstands.Onthewesternslopethesearecommonlyfilledwith   greasewood(Sarcobatusvermiculatus).Intheshadeoftheseshrubsafewspeciesofmosses,such p!  asCrossidiumandSyntrichiaarefrequent. V"!   Gypsum/saltdomesretainwaterlongerthantheflatdesertpavements.Themostnotableone "$r# inColoradoisinParadoxValley.The_sypsum_Ԁretainsrainwaterlongerthantheadjacentsandyflats, %X $ andisdenselycoveredbya_veriety_Ԁof_cruatose_Ԁlichens.Themossfloraispoorbutespecially %>!% interesting.Didymodonnevadensisoccupiesbarefacesbetweenthelichens,whileBryum &$"& nanoargenteumandSyntrichiacaninervisoccurintheshadeoflowAtriplexshrubs. ' #'   (#( @8 8 3 ThegeographyofColoradobryophytes     Therearetwowonderfulthingsaboutbryophytedistributionswhichhavenothingmuchto ~ dowiththebryophytesthemselves.ItisaboontoscientiststhattheLorddidnotdistribute d bryophytes,orplants,forthatmatter,otherplants,atrandom.IfHeorShedid,wescientistswould J haveaterribletime,becausewewouldhavetoscoureveryinchoftheplanet.Bryophytesaretied 0  verycloselytotheirmicrohabitats,whicharediscreteandcanberecognizedbyanastuteobserver.  f Furthermore,bryologicalcollectorsmoveatasnailspacecomparedtowildflowerpeople.Wemight  L walkafewmilesinaday"longerdistancesifwecloseoureyesuntilwereachthedestination  2 whereweknowwewillfindaparticularmicrohabitat.Usuallywetendtofindaninterestingspotat    theroadsideoronlyafewhundredyardsaway,andspendthedayrightthere!      Acorollarytothefirstblessing,weshouldbeverythankfulthatmanhasbuiltroadsandtrails, z  despitethefactthatwetendtoresenttheseintrusionsonpristinehabitats.Butforthesewemight `  neverfindsomeoftherarerspecies.Whenlookingatadotdistributionmapofabryophytewemust F  rememberthatthisiswherecollectionshavebeenmade.Adotmapwillgiveusageneralideaof ,|  otherlocalities,butfewareasaresowellcollectedthatonecanmakedetaileddotmapscoveringthe b smallergeographicaldivisionssuchascounties,townships,ranges,andsections.Thusitisnotvery H meaningfultodeclareabryophyterarebecausefewcollectionshavebeenmade.Fewbryologists . collectthesamecommonmossinmorethanafewcountiesinastate.Atthesametime,becausewe  havefoundararemicrohabitatdoesnotmeanthatwehaveexhaustedthepossibilitythatothersuch  placesmaystillbefoundfartherfromtheroad.    Bryophytesoftenhavemuchlargerormorewidelydisjunctdistributionpatternsthando \ floweringplants.TheSouthernRockyMountainsdrawsitsflorafromtheobviousmigrationpathway B affordedbytheimmensenorthsouthextentoftheWesternAmericanCordillera.Everydrainage (x systemthatradiatesfromthecoreofthesystemalsoservesasahighwayofmigrationforriparianand ^ lowlandordesertspecies.Anaptanalogyisthatofagreatwheel,whosehub,theSouthernRockies, D preservesthemostancientsurvivors,andwhoseaxleandspokes"theCordilleraandthedrainage * systems,providethepathwaysalongwhichmigrationhastakenplacethroughtimeunderthe  pressuresofclimaticshiftsand_orogenic_Ԁmovements.    TheColoradobryophytefloraispredominantlyborealmontane,astestifiedtobythefactthat r!  allbutaverysmallnumberofspeciesarecommontoColoradoandScandinavia.Abryologisttrained X"! inScandinaviaorSiberiawillbeverycomfortableinColoradofortherearesomanyspeciesin >#" common. $$t#   AsignificantnumberarerestrictedtoprotectedravinesintheeasternslopeoftheFront %@!% Range,wheretheypersistasrelictsofthePleistocene.Thesespeciesofeasternwoodlands(some &&"& havingextensionstothePacificNorthwest)formerlyradiatedtothefootoftheRockyMountains. ' #' Followingthedryingoutofthehighplains,theywereeliminatedfromtheinterveningterritory. (#( AmongtheseareAmphidiummougeotii,Anomodonspp.,Brachytheciumacuminatum,Cnestrum )$) schistii,Dicranumbrevifolium,D.flagellare,D.fulvum,D.montanum,Didymodontectorum, n*%* Entodoncladorrhizans,Grimmiaanomala,G.pilifera,Leskeapolycarpa,Mniumhornum,Neckera T+&+ complanata,Plagiotheciumcavifolium,P.laetum,Platygyriumrepens,Rhytidiadelphustriquetrus. :,', Ї  AfewdesertandlowlandspeciesarecommontodesertandsteppeareashereandinAsia  (Anoectangiumhandelii,Didymodonanserinocapitatus,Jaffueliobryum,Crossidium,Aloina,  Pterygoneurum).Averysmallnumberofworldwidedisjunctscanbeconsideredtobeancientrelicts | ofTertiaryorolderfloras(Bryoxiphiumnorvegicum,Gemmabryumalpinum,Leptodonsmithii,Oreas b martiana,Voitianivalis).AveryfewspeciesfollowaNorthAmericanCordilleraAndeanpath H rangingfromAlaskathroughColoradoanddowntosouthernSouthAmerica(Anacolia,Bartramia . ~ potosica,Leptopterigynandrum,Rhexophyllumsubnigrum,Syntrichiabartramii).  d   Oftheborealmontaneelement,themajorityofourspeciesbelongtotaxaofrelativelydry  0 mountainareaswithcontinentalclimate.Speciesrequiringconstantlyhighhumidity(suboceanic    species)suchasthoseoftheforestsofthePacificNorthwestareeitherabsenthereortheyoccurin    smallremnantsofwhatmighthavebeenduringthePleistocenelargerareassuitedtotheirgrowth(,   Hylocomiastrumpyrenaicum,Hylocomiumsplendens,Pleuroziumschreberi,Ptiliumcrista x  castrensis,Rhytidiadelphustriquetrus). ^  ***EndemicPacificNorthwest:Brachytheciumleibergii,Roelliaroellii... *z  @ + Somethoughtsaboutecology,evolution,andtaxonomy  F   Bryophytesareobviouslycloselytiedtoparticularsubstrates,notonlythegeneraltypesof  rock,butthefeaturesofrocks.havingtodowithexposure,seepageareas,crevices;treeandshrub  habitats:thekindsofbark,theheightfromtheground,theexposure;soiltypesandconditions:soil  typesandconditions,whetherheavy,light,alkaline,compactedorloose;thesurroundingforest v canopy.Othersmaycometomind. \   Koponen(1982b)presentsaninterestingdiscussionconcerninghowbryophyteecologymay (x giveinsighttotheirtaxonomy.Hesays Itismucheasiertofindexamplesofspecieswhichare ^ strictlylimitedtoaspecialmicrohabitatthanofspecieswhichgrowinalargevarietyofhabitats, D suchasCeratodonpurpureus.Habitatspecificityofteniscorrelatedwiththeevolutionandhence * thetaxonomyofthespecies.Thisiswhyitissoimportanttoincludehabitatinformationwith  collections.WhenIwasidentifying, siteunseenDonaldMcVeansAustralianalpinecollections,  Isometimessuggestedaname. Itcantbethat;thehabitatisallwrong!Hewasrightahundred   percentofthetime.Koponensays. Iamalsoponderingwhyweshouldnotacceptaspecificstatus r!  forataxonwhichdiffersfromitsnearestrelativeinseveralcharactersandalsohasadifferent X"! ecology?Ifeelthatthisquestionispertinenttoourconsiderationofthetaxonomicpositionofthe >#" highalpine varietyofAulacomniumpalustre.Thismossnevergrowswiththespeciesproper,its $$t# leafshapeisunique;itproducesnogemmae,itisdisjunctinhighmountainsoftheEastern  %Z $ hemisphere;itdoesnotfruit;itconcealswhatstemrhizoidsitpossesseswithintheaxilsoftheleaves; %@!% Ihavenotencountered intermediates. &&"& @: _Geobotany_Ԁofbryophytes  (#(   Becauseoftheirphysicalnature,theirexistenceaspredominantlyhaploidorganisms,andtheir p*%* closeattachmenttotheverythinlayerofsubstrateonwhichtheygrowandmustsurvive,bryophytes, V+&+ morethananyotherplants,arerecognizedasbeingassociated,ifnotdependenton,variousphysical <,', substrates,oftenhavingtodowiththeavailabilityofwaterasactualliquidorasmistordew;if  liquid,whetherthewaterisrunningornot,whetheritisaugmentedbychemicalionscomingdown  fromahillsideorbyrunofffromminetailings,etc.,hatis,whatthepHofthesubstrateis,acidor | basicorneutral.Othercharacteristicsofthesubstratesonandinwhichtheygrowareimportant. b Bryophytes,atleastmanyofthem,arecategorizedasbeing_calciphiles_Ԁor_basiphiles_Ԁ(associatedwith H calciumorothersubstancesincludingpotash,similarinchemicalbehavior);disturbanceplants, . ~ _nitrophiles_Ԁ(requiringortoleratingnitrogenousmaterial);dung lovingmosses;minerotrophic  d mosses.  J   Notverymanymossesarewellenoughknownforthemtobecategorizedwithanyofthese    terms,andmoreover,itisnotunderstoodwhattherolesofsubstratereallyplaysintheoccurrence    ofmosses.Mossesthatareconsidered_calciphiles_Ԁononepartoftheworldmaynotbehaveassuch   inanother.MossflorasinPlaceslikeScandinaviaaremorethoroughlyclassifiedthanothersbecause x  ofalonghistoryofinterestinthemandbecausecalciferousrockassociatedwithabundantmoisture, ^  makescalciferoussubstratesmoreobviousselectorsofplantspecies.IntheRockyMountains, D  however,wenotonlyhavehadlittleseriousinterestinmosses,andthesocalled_calciphiles_Ԁareoften *z  notdefinitelyconnectedtocalciferoussubstrates.VerylittlenotehasbeenmadeinColorado ` concerningthecalciumcontentofthesubstrate. F   HereisapartiallistofColoradospeciesthatareconsideredbyCrum_(1973a)_Ԁasbeing  _calciphiles_,orlikely_calciphiles_.Afewmorehavebeenaddedbythepresentwriters.Thoseindicated  as(!Crum)weconsidersurprising,sinceinColoradotheydonotnecessarilybehaveassuch.  e( X5(#(#(Abietinellaabietina(!Crum) Z Aloinarigida @ Anoectangiumaestivum &v Anoectangiumhandelii  \ Anomodonrostratus B Anomodonattenuatus ( Barbulaconvoluta  Barbulaunguiculata  Brachytheciumturgidum( probably!Crum)   Breidleriapratensis(!Crum) p!  Campyliumstellatum V"! _Catascopium_Ԁnigritum <#" Conardiacompacta "$r# Cratoneuronfilicinum %X $ Cratoneuronfilicinum %>!% Dicranellavaria &$"& Didymodonanserinocapitatus ' #' Didymodonrigidulus (#( Didymodonsubandreaeoides )$) Distichiuminclinatum l*%* Drepanocladusaduncus(!Crum) R+&+ Encalyptaprocera 8,', Grimmiaanodon Z, Gymnostomumaeruginosum @- Homomalliumadnatum &v. Hygrohypnumluridum  \/ Hymenostyliumrecurvirostrum B0 Hypnumcupressiforme(!Crum) (1 Myurellajulacea 2 Orthotrichumanomalum 3 Orthotrichumpellucidum  4 Plagiopusoederianus p!5 Pseudocalliergonturgescens V"6 Pterygoneurumovatum <#7 Rhytidiumrugosum(!Crum) "$r8 Saelaniaglaucescens %X 9 Scorpidium(allspecies) %>!: Sphagnumwarnstorfii &$"; Syntrichianorvegica(!Crum) ' #< Syntrichiaruralis(!Crum) (#= Tomentypnumnitens(!Crum) )$> Tortellatortuosa l*%? Tortellafragilis R+&@ Tortula(_Phascum_) 8,'A Tortulaobtusifolia  Weissiacontroversa   |   ( X5(@ 2Otherkindsofsubstraterelationships. H Disturbance:Ceratodon,Bryumargenteum,B.angustifolium,Gemmabryumsubapiculatum  d _Nitrophile_:Ceratodonpurpureus  0 _Basiphile_:Ditrichumflexicaule   Minerotrophic:Coscinodoncribrosus,Mielichhoferia x  Fireash:Funariahygrometrica,Leptobryum D  Dung:Splachnaceae(Tayloria,Splachnum,Voitia) ` @vv>WinterActivity ,   Weusedtothinkthatthefirstsnowwouldbringanendtoourworkinthefield,because,we  thought,oncethemossesbecomecoveredwithsnow,therewouldbenouselookingforthemuntil  MayorJune.AtHallRanchandontheoutwashcolluviumaroundBoulder,wefoundthatthisisnot t necessarilythecase. Z   In1005wehavebeenverysuccessfulinfindingmossesonwhatordinarilywouldbe &v consideredextremelydryhabitatsatlowaltitudes.However,inthewinter,snowcoverisspotty,and  \ groundmossessuchasTortulaacaulon,canbeveryabundantlocallyingrasslands,their B sporophytesperfectlyformedalthoughgreen,readytopopwhenspringcomes.OntheHallRanch, ( SyntrichiacalcicolaandGrimmialongirostrisareactiveintheshelteredgulchesundertheshrubs.  Ontherimrock,Grimmiaanodonbearsyoungcapsules,DidymodonrigidulusandD.  anserinocapitatusareactive,andintheshallowcrevicesoftherimrock,Pterygoneurumsubsessile,   Encalyptavulgaris,andWeissialigulifoliaalsoaredevelopingcapsules.Intheextremelydeep, p! narrowcreviceswherethereisa_littl_Ԁmoisture,onecanusuallyfindAnoectangiumhandelii,aswell V" asFissidens_sublimbatus._ <#    ThissituationsurelyobtainsinthecanyonsandplateausoftheWesternSlope,but %X " unfortunatelywehavenothadtheopportunitytovisittheseareasinthewinter.Aresidentbryologist %>!# needstoexplorethegypsumflatsforCrossidiumandAloina,andtherimrockforEntosthodon &$"$ sonorae,Didymodonconvoluta,Funariaamericana,andotherminuteephemeralspecies.Thereis ' #% muchworktobedonealsoonthehighplains,wherethereisalwaysapossibilityofdiscovering (#& _Aschisma_Ԁ_kansanum_Ԁontheundersideofquartzpebblesintheeasternmostcounties.Theweatheris )$' alwaysaprobleminthewinterandspring,andthemidgescanprovedaunting.  l*%( @> PARTI.MOSSES   @ 3 Keystothegeneraof_colorado_Ԁmosses  ~ (Inpreparation)@E  L @ 3Family,Genus,andSpeciesTreatments  @> AMBLYSTEGIACEAE  b Ref:Hedenspapers 0    PleurocarpousmossesinfensgenerallyaresuggestiveofAmblystegiaceae,butbelongto  L diversefamilies.Forecologistsworkinginfens,itisveryusefultobeabletoidentifytheplantsin  2 thefield.Thefollowingfieldkeywillmakethejobeasier.Note:Leaffeaturesrefertothestem    leaves,notthebranchleaves.    @  ' Fieldkeytothepleurocarpousfenmosses(severalfamilies) z   @E1a.0  Leavesbroadlyovate,ovateorroundedtriangular,inupperpartsuddenlyroundednarrowed H  orapiculate(note:theinflexedupperleafmarginsmaygivetheapexamoreamucronate .~  appearance)#@"@"x(#. #(#(##(2) d 1b.0  Leavesstraightorfalcate,fromovateortriangularbasalportiongraduallynarrowedtowards J leafapex#@"@"x(#.r r $(#(##(5) 0 2a.0  Costashort,doubleorsingle,notorhardlyvisiblewithahandlens;shootsturgid,slightly  andirregularlybranched;alarcellsnothyaline,inanindistinctlydelimitedgroup.   Pseudocalliergonturgescens x(#(# 2b.0  Costalong,reachingleafmiddleorfurther,singleorbranched,usuallyeasilyseenwitha ` handlens#@"@"x(#. !(#(##(3) F 3a.0  Leavesusuallyovateornarrowlyovate;plantsofalpinepools.#@"@"x(#.W(#(##(4) b 3b.0  Leavesusuallybroadlyovateorbroadlyroundedtriangular;greenorbrownishgreen H species,sometimeswithapalepinkishhue. Calliergon .(#(# 4a.0  Redcolorscommon,otherwisegreentodarkgreen;usuallystronglybranched;stemleaf  apexusuallydistinctlyapiculate,atleastinyoungleaves;leafpointoftenbentinwards;very   rarelyfoundwithleafbornerhizoids. Warnstorfiasarmentosa x! (#(# _4b_.0  Paleoryellowgreen,usuallysparselybranched;leafapexroundedorroundedobtuse;leaf `"! bornerhizoidscommonneartheleaftips. Straminergonstramineum F#"(#(# 5a.0  Costashort,usuallydouble,notorhardlyvisiblewithahandlens#@"@"x(#.[(#(##(6) %d $ _5b_.0  Costalong,reachingmiddleofleaforfurther,normallysingle,usuallyeasilyseenwitha %J!% handlens#@"@"x(#. !(#(##(8) &0"& 6a.0  Verylargespecies;shootsoftenturgid,notflattened;greenoroftenwithbrown,yellow (#( brown,red,orblackishcolors. Scorpidiumscorpioides )$)(#(# 6b.0  Mediumsizedspecies;shootsoftensomewhatflattened;greenorpaletoyellowgreen z*%* species#@"@"x(#.H H (#(##(7) `+&+  F,', 7a.0  Leaveswithlargeanddistinctgroupsofalarcells;costashortandforked,orlacking.leaves  stronglyfalcate Calliergonella(SeeHypnaceae) (#(# 7b.0  Leaveswithsmallandindistinctgroupsofalarcells;costaveryshortandinconspicuous; ~ leavesweaklyfalcate. Breidleriapratensis(SeeHypnaceae) d(#(# 8a.0  Leavesstraight,lanceolate,stronglyplicate#@"@"x(#.E(#(##(9) 2  8b.0  Notasabove#!!w(#.  $(#(##(10)  h 9a.0  Undersideofthestraightstemsdenselyclothedwithbrowntomentum. Tomentypnum  4(#(# 9b.0  Undersideofstemsnottomentose. Brachytheciumturgidum   (#(# 10a.0  Stemleavesusuallymoreorlessbroadlytriangulartoverybroadlycordate,ratherquickly   narrowedtotheacumen;branchleavesmuchsmaller,falcate;alargroupslarge,triangular   andwelldelimited,reachingfrommargintocosta;plantsusuallydenselypinnate;leavesnot f  plicate. Cratoneuronfilicinum L (#(# 10b.0  Leavesroundedtriangular,ovateorbroadlyovatetolinear,moregraduallynarrowedtoward 4  theapex;alargroupsusuallylessdistinct;branchingrarelydenselypinnate(tomentummay j occurinPalustriellafalcata,Tomentypnum,andConardia).#!!w(#.R(#(##(11) P 11a.0  Paraphylliapresent(tearoffafewleaves;theparaphylliaareusuallydistinctlyvisiblewith  ahandlens);largespecieswithfalcate,distinctlyplicateleaves;costastrong;tomentum  frequentlypresent. Palustriellafalcata (#(# 11b.0  Paraphylliaabsent;tomentumlackingexceptinTomentypnum#!!w(#.R(#(##(12)  12a.0  Leavesconspicuouslyplicate;commonlyfruiting,thecapsulesnormallyhorizontal(not L commoninfensbutverycommoninwetspruceforests). Sanioniauncinata 2(#(# 12b.0  Leavesnotplicateorindistinctlyso#!!w(#.((=(#(##(13) j 13a.0  Leavesfrommoreorlessstraightandrathererectbasalportionwithratherstronglycurved 6 upperpart;alargroupsundifferentiatedorverysmallandnotvisibleinthefield.#!!w(#. p(#(##(14)  13b.0  Leavescurvedmoreorlessalongtheirentirelength,ornearlystraight;alargroupsdistinctly  differentiatedandusuallyvisibleontornoffleaves(indistinctlydifferentiatedin   Pseudocalliergonangustifolium)#!!w(#.LL7(#(##(15) ~!  14a.0  Shootsrelativelystiffandmoreregularlybranched;green,yellowgreen,brown,orbrown J#" red(auniquecolorcombinationincontrasttothenext);leavesgreenoryellowgreenwith 0$# brownredcosta(oftenalsoleafbase),somewhatdull(duetorelativelyshortleafcellswith %f $ squarishends). Scorpidiumcossonii %L!%(#(# 14b.0  Shootssomewhatlargerandlessbranched;red,blackishred(brownred)orgreen;glossy &4"& (duetolongcellswithgraduallynarrowedends. Scorpidiumrevolvens '#'(#(# 15a.0  Costalongexcurrent;shootandbranchapicessometimespencillike;usuallygrowing )$) submergedinpools#!!w(#.& & *(#(##(16) ~*%* 15b.0  Costanotexcurrent"incaseofdoubt(occurringinspecieswithradiallybranchedshoots) d+&+ theshootandbranchapicesarenotpencillike;submergedornot#!!w(#.ddY(#(##(17) J,', Ї16a.0  Shootandbranchapicespencillike;leavesstraightorcurved,oftendeepredpurple;young  axillaryhairslongandbrown(visiblewithahandlensifafewleavesaretornoff);shoots  radiallybranched. Warnstorfiatrichophylla |(#(# 16b.0  Shootandbranchapicesneverpencillike;leavesusuallycurved,neverbecomingred;young d axillaryhairssmall,hyaline(notvisiblewithahandlens);shootsdistichouslybranched. J  Drepanocladuslongifolius 0 (#(# 17a.0  Yellowbrown,brownishyellow,orgreenspecies,typicallywithagoldenglosswhendry;  N alarcellsindistinctlydifferentiated;plantsofstronglycalcareoushabitats. Pseudocalliergon  4 angustifolium   (#(# 17b.0  Colorvarying,notwithgoldenglosswhendry;alargroupsusuallylargeandmoreorless    distinctlydifferentiated;inlessstronglycalcareoussites#!!w(#.T(#(##(18)   18a.0  Shootsdistichouslybranched,neverred;rhizoidsnevergrowingfromtheleaves;mostlyin f  nutrientrichhabitats#!!v(#. .(#(##(19) L  18b.0  Shootsmoreorlessradiallybranched,mostdistinctwhengrowingwithsteminvertical 2  position;sometimeswithredcoloration,andsometimeswithrhizoidsgrowingfromthe h leaves;inlessnutrientrichsituations. Warnstorfia N(#(# 19a.0  Planthabit Drepanocladuslike,thatis,withleavesfalcatesecundtosometimesstraight  anderect;leafacuminainstraightleavedplantsplaneoratmostslightlyfurrowed.   Drepanocladusaduncus (#(# 19b.0  Planthabit Campyliumlike,thatis,withleavesfromstraightanderectbasesusuallywith  leafacuminamoreorlessspreadingorsquarrose;leafacumenfurrowed. Drepanocladus f polygamus N(#(#  @' ' . Hintstorecognitionofgeneraofflowingwater  R   Aquaticpleurocarpousmossesthatgrowoverseepingrocksorattachedtorocksinsmall   streamsmaybelongtogeneraofseveralfamilies.Brachytheciumrivularemaybemistakenfor  Amblystegium,buttheleavesaremorebroadlyovateandfinelyserrulatedistally,andthealargroup   isconspicuouslyinflated. !  @@ Generalkey  N#" la.0  Alarcellsinflated,sharplydifferentiated;paraphylliausuallypresent;leafcellsmoreorless %l $ papillose(fromprotrudingcellends). Cratoneuron and Palustriella &R!%(#(# lb.0  Alarcellsvarious;paraphylliaabsent;leafcellssmoothornearlyso#!!w(#.88^(#(##(2)) &:"& 2a.0  Leavesusuallysquarrose. Campylium andrelatives (SeeCampyliaceae) ($((#(# 2b.0  Leavesneversquarrose#@"@"x(#.VV.(#(##(3) )$) 3a.0  Leaveswithasinglecostaextendingatleasttomidleaf#@"@"x(#.Q(#(##(4) j+&+ 3b.0  Leaveslackingacosta,orthecostashortanddouble#@"@"x(#.rrM(#(##(8)  4a.0  Leavescordateovate,oblongovate,oroblong,neverlongacuminate;marginsentire. |  Calliergon anditsrelativesb(#(# _4b_.0  Leavesalwaysdistinctlyacuminate;marginseitherentireorserrulate#@"@"x(#.^(#(##(5) J 5a.0  Leavesusuallyfalcate(exceptinsomesubmergedforms)orplicate,orboth. Drepanocladus  f  anditsrelatives N(#(# _5b_.0  Leavesneitherdistinctlyfalcatenorplicate#@"@"x(#.zzE(#(##(6)  4 6a.0  Costastrong,percurrentorexcurrent;plantssubmergedinrunningwater. Amblystegium  (#(# 6b.0  Costashorter;plantsterrestrialorinwetplaces,rarelysubmerged#@"@"x(#.\(#(##(7)   7a.0  Leavesverysmall,2mmorlesslong;plantssmallanddelicate;plantsofforests#@"@"x(#.4 4 j(#(##(8) d  7b.0  Leaveslarger;plantsrathercoarse;plantsofwetlands#@"@"x(#.O(#(##(9) J  8a.0  Leavesandbranchesverysmallandslender,leavesupto0.45mmlong,ovatelanceolate, f neversquarrose,entireoroftenserrulateatthebase;2-3celledgemmaepresentintheleaf L axils. Platydictya (seeHypnaceae)2(#(# 8b.0  Leaveslarger,otherwisenotasabove. Amblystegium (#(# 9a.0  Leavessmallorlarge,lessthantwiceaslongaswideor,iflonger,thensomewhatfalcate;  basalcellsrarelywithpittedwalls;onwetrocksinornearstreams. Hygrohypnum ~(#(# 9b.0  Leavesusually2mmlongorlonger,commonlytwiceaslongaswideormore,deeply f concave,neverfalcate;wallsofbasalcellsoftenpitted;notattachedtorocks,buteither L submergedinfenpondsorinloosegravelofsnowmeltrills0 (#(#  0  (# (#  #!!w(#.Z (# (##(10)  2  10a.0  Leavesmoreorlessfalcatesecund,atleastthoseofthetipsofthestemsandbranches;alar P cellshyaline,thinwalledandinflated,insmall,inconspicuousgroups. Scorpidium 6(#(# 10b.0  Leaveslooselyimbricatetospreading,notatalloratmostveryslightlysecund;alarcells  shortlyoblongand_subquadrate_,neitherhyaline,thinwalled,norinflated. Pseudocalliergon (#(# @x x 4 Amblystegium Bruch&Schimper,1853 !  1a.0  Leaveslessthan1mmlong,generallyserrulateforatleastpartoftheirlength;basal P#" marginalcellsandadjoiningalarcells_subquadrate_Ԁorshortrectangulartotransversely 6$# elongate,firmwalled. A.serpens %l $(#(# 1b.0  _Lea vesw_Ԁmorethan1mmlong,entire;basal_marinal_Ԁcellsshorttoratherlongrectangular, &:"& theadjoiningalarcellsquadratetorectangularandratherlax. A.riparium ' #'(#(#   A.riparium (Hedwig)Bruch&Schimper.Commoninverywetplaces,particularlyinareas )$) wherelivestockorotherpastoralactivityisstronglyevident.Ihavefounditabundantindrinking *%* troughs,oncheckdamsluicesandinrelativelystillwaterthroughthemiddlealtitudes. l+&+ Ї  Inourdistributionofspecimens,wemadeahorrendousmistakeofthinkingalarge,robust,  submergedpopulationinthebedofanintermittentstreamwasFontinalishypnoides.Evidentlywe  werenotalone.Crum&Anderson,p.994,write: Amblystegium_laxirete_representsanextreme | developmentofaFontinalislikehabit.Itsometimesoccursinlong,streamingmassesinswiftly b flowingwaters....Inthishabitat,theplansarestriking,butinlessvigorouslyflowingwatersthey H arelessdistinctive.Anumberofothernamedformsevidentlyarevariantsofthisextraordinary . ~ modification.Moral:DontassumethateverythingthatlookslikeFontinalis,isFontinalis!  d   A.serpens (Hedwig)Bruch&Schimpervar. juratzkanum (Schimper)Rau&_Hervey_.A  0 small,nondescriptspecies,usuallyfruiting.Thespeciesisgenerallydistributedonwetboulders,soil,    basesofsaplingsorshrubs,exposedroots,rottenlogs,etc.,fromthefoothillsuptothesubalpine.    Theleaveswidespreading,about0.6mmlongandgracefullyacuminate,withcostatoaboutmid   leaf.Themedianleafcellsareabout6:1or35mx5-7m,smoothandmoderatelythickwalled. z  Theleafmarginisslightlydenticulatefromprojectingdistalendsofthecells.Thealarregionis `  quiteclearlydifferentiated,consistingofquadrateandbroadercellsformingamoreorlesstriangular F  patchatthebasalangles.Thecapsuleiscurved,theoperculumconic,andtheurnstrongly ,|  constrictedbelowthemouth.WeareinclinedtofollowNyholmintreatingA.juratzkanumasa b varietyofA.serpens.Thevarietyisdistinguishedbyhavingtheleavesmorewidelyspreadingand H withthemarginalbasalcellsrectangularinsteadofquadrate. .   A.varium (Hedwig)Lindberg.Anexceedinglyvariablespeciesofrunningwaterinthe  loweraltitudes,havinghadmanyformsrecognizedasseparatealthoughindistinctspecies,place  eitherinAmblystegiumorHygroamblystegium.CrumandAnderson(1981)amplyshowthe x frustrationsinherentuponseparatingthem.Hygroamblystegiumhasbeenaterribleproblembecause ^ ofitsvariability.Recently.molecularbiologyhascometotherescue(see_Vanderpoorten_Ԁ2004),in D whichtheauthordemonstratesthatH.fluviatile,H.humile,H.noterophilum,andH.tenaxare *z justifiably_synonymized_ԀunderAmblystegiumvarium. ` @  - Calliergon (SullivantinA.Gray)Kindberg,1894 ,   Calliergonisgenerallynotaplantoftheedgesofswiftlyflowingbrooks.C.richardsonii  growssubmergedinstillpools.C.cordifoliumoccursonsaturated,swampygroundinforest   clearingsfilledwithtallwillows,C.giganteumwasfoundonslopingrockfacesbesideaquiet v!  backwateronalevelbenchofamountainstream.AfewspeciesformerlyincludedinCalliergonare \"! commoninalpinepools.TheseincludeWarnstorfiasarmentosaandStraminergonstramineum. B#" 1a.0  Costausuallyendingwellbelowtheleafapex,withshortbranchesorforkedattheapex; %^ $ shootswithratherlong,thickbranches,moresparselybranchedandwithbranchleavesmore %D!% erectorimbricatethaninC.giganteum(seebelow). C.richardsonii &*"&(#(# 1b.0  Costaendingalmostintheleafapex(appearingtoreachtheapexasseenwithahandlens) '#' #@"@"x(#.(#(##(2) (#( 2a.0  Costastrong;alargroupsofstemleaveslarge,triangularandsharplydelimitedfrom t*%* surroundingcells,extendingfromleafmargintooralmosttothecosta;leavesbroadly Z+&+ triangular(shootswhenwelldevelopeddenselybranchedlikeasprucetree,withmoreor @,', lessspreadingbranchleavesexceptnearbranchapices. C.giganteum (#(# 2b.0  Costaweaker;alargroupssimilarbutdiffuselylimitedfromsurroundingcells;rarelyfound  inpermanentlysubmergedhabitats). C.cordifolium ~(#(#   C.cordifolium (Hedwig)Kindberg.Poolsandlakesideswamps,uppermontaneand L subalpine.OurrecordsarefromtheplateausofwesternColoradoandfensintheFrontRange. 4    C.giganteum (Schimper)Kindberg.Wehavetwocollections:LarimerCo.:Cirque  P Meadowstrail,vic._Pingree_ԀPark,streamlettributaryofFallCreek,9,600ft.,Hermann&Rolston  8 80114.Samelocality,Weber&Wittmann112731.      C.richardsonii (Mitten)Kindberg.Submergedinfens,Boulder,Larimer,andClearCreek   counties.TheBoulderCountyspecimensareextremelylargeleaved,resemblingC._megalophyllum_   (soreportedbyWeber,1973). h  @ 3 Cratoneuron (Sullivant)Spruce,1867 4    C.filicinum (Hedwig)Spruce.Thisspeciesisabundantinwetsitesinthesubalpineforests R andwillowcarrsandfens.Theplantsaresmall,pinnatelybranched,andthestemleavesare : conspicuouslylargerthanthebranchleaves.Thebranchleavesareusuallyfalcate.Acharacteristic   featurearetheparaphyllia,ofvariousshapesandsizes,minutely_leaflike_,whichunfortunatelyvary  inabundanceandaresometimesalmostlacking.Thestemleavesarebroadlyovate,_witrh_Ԁavery  strongcosta.Themedianlaminalcellsarenarrowlyrhomboid,anthealarcellsareenlarged,ina  conspicuoustriangulargroup.Thespeciesissaidtobestrongly_calciphilous_,andiscommonin h calcareousfensinthesubalpine. N @d d 2 Drepanocladus (MllerHal.)Roth,1899 j   CrumandAndersonacceptDrepanocladusandScorpidiuminatraditionalsense,whichfor 8 fieldbotanistsseemsreasonable,andtheyrejecttheseparationofseveralotherspeciesintothe  generaLimprichtia,Warnstorfia,andScorpidium.However,wefollowtherecentworksofHedens.    HedenssuggestsonewaytodistinguishDrepanocladusfromWarnstorfia: Itisalways !  goodtolookforrhizoidalinitialsneartheleafapices.Ifyoustudy1015leaves(orashootapexwith f"! severalleavesleft)inthemicroscope,therearealmostalwaysatleastsomeleaveswithsuchinitials L#" inWarnstorfiaspecies,butneverinDrepanocladuss.str. 2$# 1a.0  Leavesfrommoreorlessstraightanderectbases,usuallywiththeacumenmoreorless %N!% spreadingorsquarrose;leafacumenfurrowed;plantsmoregoldenbrownthangreen. D. &4"& polygamus '#'(#(# 1b.0  Leavesstronglyfalcatesecund;leafacumeninlessfalcateleavedplantsplaneoralmost ($( slightlyfurrowed;plantsgreenwithnoothertints#@"@"x(#.K(#(##(2) )$) _ 2a.0  Costaofstemleavesendingwellbelowtheleafapex;leafmarginentire,oronly f+&+ occasionallyveryfinelydenticulate. D.aduncus (#(#  2b.0  Costaofstemleavesexcurrent,rarelyendingafewcellsbelowleafapex;oneorbothleaf  marginsusuallypartlyfinelydenticulate. D.longifolius ~(#(#    Drepanocladusaduncus (Hedwig)Warnstorf.Anextremelycommonandvariablespecies, L verywideranginginaltitudefromtheplainsuptothealpine.Theelongate,inflatedalarcellsina 4  rowcontinuingalmosttothecosta,isdiagnostic.Theplantisweak,notstandingerect,yellowish  j green,neverwithreddishorbrowncolors,verylittlebranchedandthennotatallpinnate;thecosta  P isslenderandelongate.However,somecollectorshaveconfusedthiswithCalliergonella(was  6 Hypnum)lindbergii.Thelatterstandsstifflyerectincloseorder,theleavesareverybroadatthebase    (triangularovate)andwithlargealarcellsatthebasalanglesjustwheretheleafbecomesdecurrent.    Thecostaisabsentorshortanddouble,andfaint,andtheleafmarginstendtocurveinwardnearthe   apex. ~    Drepanocladuslongifolius (Mitten)Paris.Insubalpinepools.Wehaveoneexcellent J  collection:GrandCo.:RockyMt.Nat.Park:FloatingonpondalongUteTrailsouthofLakeIrene, 2  10,400ft.,Hermann26506.Hedens(_BFNA_)acceptsthisforColorado,saying: D.longifolius[D. h _capillifolius_ofauthors]differsfromallotherAmericanDrepanocladusspeciesinitsexcurrentleaf N costa.BecauseofthelatteritcouldhardlybeconfusedwithanyearlierDrepanocladuss.l.inNorth 4 AmericaexceptWarnstorfiatrichophylla.However,theleavesarealwaysgreen,thereareneverany  rhizoidalinitials,andtheshootsaredistichouslypinnatelybranched.InWarnstorfiatrichophyllathe  shootsareradiallybranched,andtheshootsandbranchapicespencillike,theleafmarginsaremore  stronglydenticulatethaninD.longifolius;itfrequentlybecomesredwhenemergentwhereasD. | longifoliusnevergetsred,andtheaxillaryhairsconsistof17earlybrownuppercells(12elongate b hyalinecellsinD.longifolius). H   Drepanocladuspolygamus (Bruch&Schimper)Hedens.Onerecord,fromasubalpinefen d onDiamondLake,intheFrontRange,collectedbyHermann(!Lawton).SeeFlowers,plate119:9 L 11.(Campyliadelphus,Campylium,Drepanocladus).Thishasuntilrecentlybeenplacedin 2 Campyliumandindeedhasmoretheaspectofthatgenus.Theplantisdefinitelystouterandthe  leaveslessfalcatethanD.aduncusandhasmorebrownishtints.D.polygamousisautoicous,while  D.aduncusisdioicousbutthisisdifficulttodeterminewithoutcapsules.   @8 Hygrohypnum Lindberg,1872 `"! 0   Ref.:Jamieson(1986a).$~#(#(# ***AddH.styriacum(Limpricht)BrotherusCheckalso(11).Haveweleftoutsomething? %d $ 1a.0  Stemcrosssectionwithanepidermisofenlarged,fragile,thinwalledcells,theinnercortical &0"& cellsin34layers,thickwalled,thecorecellslargeandthinwalled. H.ochraceum '#'(#(# 1b.0  Stemcrosssectionwithanepidermisofseverallayersofsmall,thickwalledcells,theouter (#( layernotfragilenorthinwalled#@"@"x(#.9(#(##(2) )$)