WPC( u'%DiF],8ja8,EM_}ž"}??x p-AI2C4f>Jk3-[b"9&DR鱓UVW?tS@|E!Con-Fgk* *Lcb! {G}u>Fh.uZp,yؑ61 y[egnr8z:9eFϏ!A ! !ֽZ;j7@* 0 N* 0* 0+ 0, 0- 0. 0w/ 0V0 021 0 2 02 03 04 0l5 0?6 07 07 08 0m9 01: 0 ; 0; 0< 0= 0Z> 0&? 0? 0@ 0xA 0pB 0hC 0ZD 0lXE 0nE 0t2F 0nF 0tG 0tG 0tG 0tpH 0tH 0tXIU6IU*J 0+C,JU>oJUNJ AJK 0D`L 0KL D-LMM)NN 0(DZOO$PJP 1QQ 72R BCR D5`R D5R BR D5R BS D59S BnS D5S BS D1S D5T B*CT D5mT B*T D5T B*U D5+U B*`U D5U B7U B7U-VVKWWkXXYYY#ZZA[A[A[[b\\ 04]]O^^j_j_j__`aa9bbYccsddddeff2g2g2ggMhhjijjjj!kkkk=ll]m]m]mmtnnnno ppGq]qqqqqqyryryryryryrssssss(ttttttJuuuuuudvvvvvvwxxxxxx1y1y1y1y1yyOzOzOzOzOzzl{l{l{l{l{||||||&}}}}}}B~~~~~~~~\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\\J%\  `$Times NewRomanX  3|I%\  `$Times NewRomanXXj\  P6G;XP($fv1(hp color LaserJet 2550 PCL60F1 \U flabelliformfimbriatumlingulategirgensohniirussowiiminerotrophicwarnstorfiifuscumhummockySandbeachutriculataresorbedflexuoseplatyphyllumSubsecundacommissuralrecurvumBeauvantheridial inflorescencessquarrosumterescorrespSphagnaceaeEvansiaB.McQueensquarroseSphagnaHattori CrumcontortumSPLACHNACEAESplachnaceaeFunariaVoitiaTayloriaSplachnumperistomeTetraplodonmnioidessphaericumlingulataexsertedcolumellahornschuchiifroelichianaGunnisonHuerfanoAbietinellaAnomodon(9 Z6Times New Roman Regular3#37=CIQYag1.a.i.(1)(a)(i)1)a)(;3$2#  0  .3  0  (5hCEKQW]cioAutoList11.1.1.1.1.1.1.1.i)http://www.nybg.org(O;$0  2#  a  .3  0` (#(#S37;CO_s11.01.0.01.0.0.01.0.0.0.01.0.0.0.0.01.0.0.0.0.0.01.0.0.0.0.0.0.01.0.0.0.0.0.0.0.0(;3$2#  0  .3  0  (O$2G+J 0_level1  X /%4 4 <DL/23  ..  2( 4 <DL2  2D+J 0_level2   ," <DL,23  ..  2( 4 <DL2  2A+J 0_level3   ) <DL)23  ..  2( 4 <DL2  2>+J 0_level4  ` &<<DL&23  ..  2( 4 <DL2  2;+J 0_level5   #DL#23  ..  2( 4 <DL2  28+J 0_level6    DL 23  ..  2( 4 <DL2  25+J 0_level7  h DDL23  ..  2( 4 <DL2  22+J 0_level8   L23  ..  2( 4 <DL2  2/+J 0_level9    L23  ..  2( 4 <DL2  2GJ 0_levsl1  X /%4 4 <DL/23  Ԁ  2( 4 <DL2  2DJ 0_levsl2   ," <DL,23  Ԁ  2( 4 <DL2  2AJ 0_levsl3   ) <DL)23  Ԁ  2( 4 <DL2  2>J 0_levsl4  ` &<<DL&23  Ԁ  2( 4 <DL2  2;J 0_levsl5   #DL#23  Ԁ  2( 4 <DL2  28J 0_levsl6    DL 23  Ԁ  2( 4 <DL2  25J 0_levsl7  h DDL23  Ԁ  2( 4 <DL2  22J 0_levsl8   L23  Ԁ  2( 4 <DL2  2/J 0_levsl9    L23  Ԁ  2( 4 <DL2  2GJ 0_levnl1  X /%4 4 <DL/23   2( 4 <DL2  2DJ 0_levnl2   ," <DL,23   2( 4 <DL2  2AJ 0_levnl3   ) <DL)23   2( 4 <DL2  2>J 0_levnl4  ` &<<DL&23   2( 4 <DL2  2;J 0_levnl5   #DL#23   2( 4 <DL2  28J 0_levnl6    DL 23   2( 4 <DL2  25J 0_levnl7  h DDL23   2( 4 <DL2  22J 0_levnl8   L23   2( 4 <DL2  2/J 0_levnl9    L23   2( 4 <DL2  <dX:HTML PretagKD<6X9`(Courier NewKXXX?E%2A`Arial?<dX:HTML TeletypKD<6X9`(Courier NewKXXX?E%2A`Arial?6dX4HTML CodeKD<6X9`(Courier NewKXXX?E%2A`Arial?<g[:HTML Code DeKD<6X9`(Courier NewKXXX?E%2A`Arial?42HTML VarXXXXXX<:HTML BlockquXXXXXX<:HTML AddressXXXXXX64HTML CiteXXXXXX<:HTML Headin5  XXX<:HTML Headin4  XXX<:HTML Headin3  XXX<:HTML Headin2 XXX XXX<:HTML Headin1  XXX<:HTML Heading  XXX<6X9`(Courier New%2A`Arial(2G$ !  %\  `&Times New Roman(hH  Z 6Times New Roman RegularJKLM I&mage <=8C HKKKKLevel 1Level 2Level 3Level 4Level 5(3$ !  ( $ Figure  1  )!dxdx ~ FG'  _%XXAmblystegiaceae"  24   { FG'  _%XXAndreaeaceae"  38   R(  _j(#%XX  38  ڀ"Andreaeaceae R(  _..g(#%XX  24  ڀ"Amblystegiaceae(3$ !   | FG'  _%XXEncalyptaceae"  97   R(  _i(#%XX  97  ڀ"EncalyptaceaeINDEX AND CATALOG OF COLORADO MOSSES(EhCEKQW]cioAutoList21.1.1.1.1.1.1.1. z FG'  _%XXEndMatter"  248  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"  99   R(  _i(#%XX  99  ڀ"Entodontaceae } FG'  _%XXAulacomniaceae"  39   R(  _>>h(#%XX  39  ڀ"Aulacomniaceae ~ FG'  _%XXLeptodontaceae"  142   R(  _g(#%XX  142  ڀ"LeptodontaceaeGrimmia | FG'  _%XXBartramiaceae"  41   R(  _i(#%XX  41  ڀ"Bartramiaceae | FG'  _%XXFabroniaceae"  100   R(  _i(#%XX  100  ڀ"Fabroniaceae  FG'  _%XXBrachytheciaceae"  45   R(  _f(#%XX  45  ڀ"Brachytheciaceae R(  _HHH(#%XX  248  ڀ"EndMatter&c$""0  ~ FG'  _%XXFissidentaceae"  101   R(  _\\g(#%XX  101  ڀ"Fissidentaceae w FG'  _%XXBryaceae"  52   R(  _""n(#%XX  52  ڀ"Bryaceae { FG'  _%XXFunariaceae"  105   R(  _j(#%XX  105  ڀ"Funariaceae } FG'  _%XXBryoxiphiaceae"  76   R(  _h(#%XX  76  ڀ"Bryoxiphiaceae } FG'  _%XXFontinalaceae"  103   R(  _~~h(#%XX  103  ڀ"Fontinalaceae z FG'  _%XXLeskeaceae"  143   R(  _DDk(#%XX  143  ڀ"Leskeaceae | FG'  _%XXBuxbaumiaceae"  77   R(  _lli(#%XX  77  ڀ"Buxbaumiaceae { FG'  _%XXGrimmiaceae"  108   R(  _j(#%XX  108  ڀ"Grimmiaceae { FG'  _%XXCampyliaceae"  78   R(  _j(#%XX  78  ڀ"Campyliaceae z FG'  _%XXMeesiaceae"  148   R(  _22k(#%XX  148  ڀ"Meesiaceae } FG'  _%XXCatoscopiaceae"  81   R(  _nnh(#%XX  81  ڀ"Catoscopiaceae | FG'  _%XXHedwigiaceae"  133   R(  _i(#%XX  133  ڀ"Hedwigiaceae { FG'  _%XXClimaciaceae"  82   R(  _j(#%XX  82  ڀ"Climaciaceae } FG'  _%XXHylocomiaceae"  134   R(  _66h(#%XX  134  ڀ"Hylocomiaceae z FG'  _%XXDicranaceae"  83   FG'  _%XXMosses,IndexandCatalog"  211   x FG'  _%XXMniaceae"  150   R(  _m(#%XX  150  ڀ"Mniaceae y FG'  _%XXHypnaceae"  136   R(  _TTl(#%XX  136  ڀ"Hypnaceae R(  _**\(#%XX  211  ڀ"Mosses,IndexandCatalog  k  { FG'  _%XXNeckeraceae"  154   R(  _j(#%XX  154  ڀ"Neckeraceae  FG'  _%XXOrthotrichaceae"  156   FG'  _%XXPlagiotheciaceae"  160   R(  _e(#%XX  160  ڀ"Plagiotheciaceae ~ FG'  _%XXPolytrichaceae"  161   R(  _RRg(#%XX  161  ڀ"Polytrichaceae z FG'  _%XXPottiaceae"  166   R(  _zzk(#%XX  166  ڀ"Pottiaceae FG'  _%XXPterigynandraceae"  193   R(  _DDd(#%XX  193  ڀ"Pterigynandraceae | FG'  _%XXRhytidiaceae"  194   R(  _i(#%XX  194  ڀ"Rhytidiaceae } FG'  _%XXSeligeriaceae"  195   R(  _h(#%XX  195  ڀ"Seligeriaceae { FG'  _%XXSphagnaceae"  197   R(  _j(#%XX  197  ڀ"Sphagnaceae | FG'  _%XXSplachnaceae"  202   R(  _i(#%XX  202  ڀ"Splachnaceae ~ FG'  _%XXTetraphidaceae"  205   R(  _,,g(#%XX  205  ڀ"Tetraphidaceae { FG'  _%XXThuidiaceae"  207   R(  _  j(#%XX  207  ڀ"Thuidiaceae z FG'  _%XXTimmiaceae"  209   R(  _k(#%XX  209  ڀ"Timmiaceae x FG'  _%XXHepatics"  211  INDEX AND CATALOG OF COLORADO MOSSES !  _  o[X` hp x (#%'0*,.8135@8:<H?AXo8n5Xdd8.J:C:\WAW\CRYPTMS\Mosses2004.wpdJC:\WAW\CRYPTMS\Mosses2004.wpd/    Revised:  December3,2004!     @ 4(AWORKINPROGRES_S_Ԁ)  b %XXe    @ 4 GUIDETOTHEBRYOPHYTESOFCOLORADO  0  @{ { &Keysandecologicalnotesbasedonfieldandherbariumstudies   @//<BYWILLIAMA.WEBER 0  @^^2FellowoftheLinneanSocietyofLondon  @R R .ProfessorandCuratorEmeritusoftheHerbarium p @OO2UniversityofColoradoMuseum,Boulder 0 @((<RONALDC.WITTMANN p @4Associate,HerbariumCOLO,Boulder @  (aWorkingDocument)  p @;November2004version  @ Xn5X%8n5WXdXd8@= TABLEOFCONTENTS    Note:thepaginationchangesaswemakerevisions.Thefamiliesarealphabeticallyarranged. ~ For generathathavebeenmovedtootherfamilies,consulttheIndexandCatalog.Portionsofthe f manuscriptthatareinneedofamplificationareindicatedby***. N Preface(#(#{(#.  j Introduction(#(#{(#.AA  6 @>PartI.Mosses    Keytothegenera(#(#{(#.  ~  Family,genusandspeciestreatments(#(#{(#.  3 J  4U eO  5  IndexandcatalogofColoradoMosses6rO  7 k(#(#{(#.kk3 f Mossspecies(alphabeticalbyspecificepithet)(#(#{(#.> 2 Derivationsofthegenericnames(#(#{(#./  Meaningsofthespecificepithets(#(#{(#.0 z Glossaryofbryologicalterms(#(#{(#. , F AuthorsofBryophytetaxa(#(#{(#. ( b 4  eO  5  6( O<   7 ej Appendices    1.LeoLesquereux(1884)(#(#{(#./     2.TheColoradoCollectionsofA.J.Grout(1914)(#(#{(#.__H \"!   3.ThetypecollectionsofT.S.Brandegee(#(#{(#.A ($x#   4.ColoradoCollectionsreportedbyT.P.JamesinWheelerExpeditionReport,1878(#(#{(#.G!G!i@(#(#{(# %D!%   5.F.J.Hermann.MossesofRockyMountainNationalPark(#(#{(#.P '#'   .Additionspaper,1976(#(#{(#.B )$)   6.OriginaldescriptionsofmossesdescribedfromColorado(#(#{(#.aaQ X+&+  >,',   7.SevilleFlowersMossesofUtahandTheWest(#(#{(#.GGF    8.Phytogeographicalcomparisons(#(#{(#.7 |   9.ColoradoBryologicalHotSpots H     BoulderMountainPark  d     MountEvans  0     HighCreekFen    Acknowledgments(#(#{(#.? ? @(#(#{(# x  Bibliography(#(#{(#.  D  (#(#{(#@?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).    8n5XXdWXd8 =n5WXdd=@B PREFACE     IfirstlearnedaboutmosseswhenIwasabouttwelveyearsold.WhenIwasinHighSchool ~ Ibuiltasmallterrariumwithmossesandlichensinit,andentereditinacontest(thiswasbeforethe d daysofScienceFairs)attheAmericanMuseumofNaturalHistory.Withthe$25.00Iwonasfirst J prize,IwasurgedbymybiologyteachertobuyasmallbookbyA.J.Grout,MosseswithaHand 0  Lens.Thus,IacquiredmyfirstmosstextbookonChristmasDay,1934,andstillfinditusefulover  f fiftyyearslater!  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)Flowers,a   longtimeprofessorofallsortsofbiologyandbotanyattheUniversityofUtah.BillFlowers,over r!  theyears,emulatedGroutinproducingmimeographedoutlinesofkeystomosses,liverworts,and X"! ferns,ofUtah.Heamassedalargecollectionaswell,andpainstakinglyproducedmarvelouspenand >#" inkillustrationsforabookonthemossesofUtah.Hedidnotlivetoseethispublished,butafterhis $$t# Аdeath,HowardCrum,bryologistattheUniversityofMichigan,editeditanditwaspublishedunder  %Z $ thenameMosses:UtahandtheWest.ForColoradobryologiststhisbookhasexcellentdescriptions %@!% andillustrationsofnearlyallofourColoradospecies.Thebookalsohasoneofthebestintroductions &&"& tomossmorphologyeverwritteninAmerica. ' #'   Therearenowseveralexcellentmossflorasavailable,butfewofthemareespeciallypertinent )$) totheRockyMountainregion.ThePacificNorthwestiswellservedby_Elva_ԀLawton'sMossFlora n*%* ofthePacificNorthwest;CrumandAndersonhaveatwovolumeMossesofEasternNorthAmerica, T+&+ andWilliamD.ReesehasrecentlypublishedaMossesoftheGulfSouth.Sharp,Crum,and_Eckel_ :,', havepublishedatwovolumeMossFloraofMexico.Ifanyoneislookingfornewworldstoconquer,  letthemplanaMossFloraofArizonaandNewMexico!    ThisguidetotheColoradomossesispatternedaftertheexcellentseriesoflittlebookswritten b bythelateProf.HelmutGamsofInnsbruck,Austria,entitled_Kleine_Ԁ_Kryptogamenflora_Ԁvon_Mittel H europa_.Theyare Excursionfloras"fieldguidestomosses,lichens,ferns,fungiandalgaeofCen . ~ tralEurope,andtheirauthorwasoneofthekeenestfieldmeninEurope.Irememberfollowinghim  d inthefieldandlisteningandlookingashepointedout(sottovocetohimself)allofthevarious  J specieshesawashepassedby.Thisisonlyoneofthemanywaysonelearnsthelowerplants.  0   Asacompromiseowingtothedesirabilityofhavinganeasilycarriedfieldmanual,an    ExcursionFloracontainsonlykeysandecologicalnotes.Formoredetaileddescriptionsandillustra   tionstheypresumetheavailabilityofmorecomprehensiveliterature.Theyalsoexpectacertain x  amountofgeneralbiologybackgroundonthepartoftheuser. ^    Thissecondeditionstilldoesnotaccomplishmyoriginalobjectiveofeventuallyproviding *z  aninstructionalintroduction,butavailablebooksandtheplannedMossFloraofNorthAmericafill ` thatneed.MuchremainstobedoneinColoradoinrefiningourconceptsofhabitatsanddistribution. F Thefirsteditionspawnedseverallocalamateurswhonowareadeptinrecognizingmosses,and , perhapsmorewillbeencouragedtoenterthefieldwiththisrevision.    Onewhoconstructsdiagnostickeyscanusuallymakeakeythatsatisfieshimself;thatis,he  candistinguishhisspeciesunderthebestofcircumstances,whenallofthenecessarythings(includ t ingsporophytes)arepresent.Itisquiteanotherthingtoproduceakeythatworksforhisstudents,and Z thathasto workevenifvitalpartsaremissing.Thisisthegreatproblemwithmosses,especially @ inanaridarealikethisone"thatmostofthespeciesarerarelyorneverfoundinfruitingcondition. &v Thus,the_sporophytic_Ԁinformation,whichoftenmakesidentificationsurerandmucheasier,cannot  \ beincludedinakey.Thewritingofthekeythusbecomesadifficultart,withinevitablyequivocal B results.Furthermore,withoutsporophytesitisimpossibletowriteasatisfactorykeytothelarger ( groups"familiesandgenera"sothatadifferentmethodhastobetried.Mrs.PatriciaNelson,my  student,workedoutaveryuseful_polyclave_ԀkeyonIBMcardscalledRandomAccessKeytothe  GeneraofColoradoMosses,whichIconsidertobeanindispensableadjuncttotheguide.   Unfortunatelytherearenocopiesleft! p!    AllofthespecieslistedintheGuidearedocumentedbyherbariumspecimensinthe <#" herbarium(COLO)oftheUniversityofColoradoMuseum,whichnowcontainsover112,000 "$r# specimensofbryophytes.Iwelcomecommentsandcriticism,andamalwaysinterestedinaddingto %X $ thecollectionsthroughcontributionsfromcollectors. %>!%   Althoughthisdistillationrepresentsmorethanfiftyyearsofsporadicresearch,theworkis ' #' nowonlywellbeguninearnest.Thestonesarelaid,butthebuildinghasnotassumeditsfinalshape. (#( Wethinkweknowwhatgrowshere,buttherewillbeadditionsassoonasactivefieldworkbymany )$) studentsbegins.Wehavesomenotionsofhabitats,buttheyareprimitiveandneedrefinement.Every l*%* observercomestothefieldwithadifferentpairofeyesandadifferentcoreofexperience,andthere R+&+ havebeentoofewofusuptonow.Ishallbesatisfiedifthisguidewillservetobringbryologywithin 8,', thecompetenceofstudentsandecologicalresearchers,andserveasaguidetovisitingbryologists  fromabroad,untilsuchtimeasamoresophisticatedworkismadepossible.    Thismanuscriptisaguidebookinprogress.Wearemakingitavailableonthiswebsiteto b makethesectionspartsavailableastheynearfinalform.Manyofthesmallerfamiliesarereadyfor H use,butsomeofthelargerormoredifficultonesarebeingworkedoncontinuously.Itishopedthat . ~ thisversionmightbeusefultoamateurs,students,_amd_Ԁprofessionalecologists.Thismanuscriptis  d anidentificationguidewithmoredetaileddescriptionsthanareusuallygiveninkeys,supplemented  J bynotesonfieldaspect,ecologicalniche,andotherinterestingobservations.Acollaborative  0 productioncontributedbyahostofprofessionalbryologistsisinpreparation,whichwillprovide    detaileddescriptionswithillustrationsofallthespecies.Tentativetreatmentsareavailableonthe    NewYorkBotanicalGardenwebsite<4 O  5  _www.nybg.org/bsci/bfna>.65?I?   7 !w?     ^  May28,2003@= 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,and_Eckel_,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       @ -ThehistoricalcollectorsofColoradobryophytes     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)of_mosses :,', 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,_JoAnn_ԀW.(***)StudentatUniv.ofColorado,collectedandpublishedontheinventoryand *z  ecologyofalpinelichensandbryophytesonNiwotRidge,BoulderCounty(Flock1978).Also ` madeinventoriesofthemossesandlichensofCapePrinceofWales,Alaska.F(#(# 0  e _Goodding_,LeslieN.(18801967),Wyomingbotanist,studentof_Aven_ԀNelson,collectedafew_moses_  intheParkRange.(#(# 0  e Grout,AbelJoel,(18671947).GroutmadeashortvisittoColoradoin1916,andcollectedinthe t vicinityoftheUniversityofColoradossummercampatTolland,GilpinCounty.See Z appendix,andSteere(1948).@(#(# 0  e Haines,Mrs.MaryParry(18261884).Anaturalistofeclectictastes,custodianofthePaleontology  \ Dept.ofamuseuminRichmond,Indiana,andanamateurhorticulturist.Althoughshenever B visitedColoradoshewasresponsibleforfunnelingseveraloddspecimens[cf.Orthotrichum ( _hainesiae_)collectedbyherfriends,Mrs.E.J.SpenceandT.S.Brandegee,tobryological  specialists(Flowers1942).(#(# 0  e Hall,Elihu(18201882).AfarmerandamateurbotanistinAthens,Illinois.CollectedwithParryin p!  Coloradoin1862.AlsocollectedplantsinTexasandOregon.V"!(#(# 0  e Hermann.FrederickJ.(19061987),receivedhisdoctorateattheUniversityofMichigan.Hiscareer "$r# wasspentattheUnitedStatesForestServiceHerbarium,wherehisspecializationwasthe %X $ genusCarexandVicia.HeretiredtoFortCollins,Colorado,wherehecollectedafineseries %>!% ofbryophytes,especiallyfromtheFrontRangeandRockyMountainNationalPark.His &$"& careercollectionsareattheUniversityofMichigan,withmanyduplicatesattheUniversity ' #' ofColorado.SeeVoss&Reznicek(1988).(#((#(# 0  e Holmen,Kjeld(***).DanishbryologistwhoiscreditedwithdiscoveringOreasmartianaonMount l*%* Evans.R+&+(#(#  8,', 0  e Holzinger,J.M.(18531929),GermanbornMinnesotan,collectedbryophytesinColoradoin  ArapahoandPikeNationalForestsin1896,andhisColoradocollectionscontributedtohis  publishedexsiccati,Musci_Acrocarpi_Ԁ_Boreali_Ԅ_Americani_.|(#(# Holzinger,Marie.CollectedinBoulderCountyin1892. H 0  e Kiener,WalterB.(18941959),SwissbornColoradomountaineer,collectedonLongsPeak(1938),  d wherehewasaclimberandguide.Hisbryophyteandlichencollectionswereacquiredbythe  J UniversityofNebraskain1960.KienerwasinvolvedinawinterclimbofLongsPeakon  0 Jan.1012,1925,inwhichhiscompanion,Agnes_Vaille_,diedinafall.  (#(# 0  e _Goodding_,L.N.(18801967)collectedinRouttCounty(Ptychostomumpallescens,No.1781   [COLO])inAugust,1903._Goodding_smaincollectionswereattheRockyMountain x  Herbarium,whosebryophytesweregiventotheUniversityofTennessee,wheretheywere ^  laterdestroyedinafirethatdestroyedtheherbarium.D (#(# 0  e Jamieson,David,contemporarybryologist,professoratFortLewisCollege,Durango,andspecialist ` inthemossfloraofthesouthwesternSanJuanMountains.AstudentofWilfredSchofield, F Univ.ofBritishColumbia,andmonographeroftheAmericanspeciesofHygrohypnum.,(#(# 0  e Lehr,Paula,contemporaryamateurbotanist,collectedbryophytesintheGunnisonBasinandElk  Mountains,andstimulatedinterestinmossesamongamateursandForestServicestaffinthe  localarea.t(#(# 0  e Nelson,LawrenceT.(18621932).Primarilyamycologist,collectedforthe_U.S.D.A._Ԁasaforest @ agentoccupiedinlocatingpoisonousplantsinColoradoandelsewhere.Hecollectedafew &v mossesfromtheGunnisonarea.Nelsontaughtatanextraordinarynumberofcollegesand  \ universitiesinmanystates.Hisbryophytecollections(19021910)areatDukeUniversity. B SeeGier(1953).((#(# 0  e Nelson,Patricia(1940),_contemporary_Ԁamateurbryologist,LibrarianattheUniversityofColorado  MedicalSchoolinDenver,studiedthemossesofClearCreekCanyon,andforhermasters   thesispreparedaveryusefulRandomAccessKeytotheGeneraofColoradoMosses.An p!  amateurbryologist,sheiscreditedwiththediscoveryoftheonlycollectionofLeptodon V"! smithiiinNorthAmerica.<#"(#(# 0  e Parry,CharlesChristopher(18231890). Duringthatyear[1865]RockyMountainmosseswere %X $ receivedfromDr.Parry,whoafewyearsbeforehadnamedColoradosRockyMountain %>!% peaksforDr.GrayandDr.Torrey)(Rodgers1940).OnlyBartramiastrictawasspecifically &$"& attributedtoParrybyLesquereux&James(1884).SeeWeber(1997).' #'(#(# 0  e Porter,ThomasConrad(18221901).Botanistandclergyman,wasattachedtotheHaydenSurveys )$) from18691874((Foster1994).Hismosscollectionswerefewandincidentaltovascular l*%* plants.ProfessoratLafayetteCollege,Lancaster,Pa.Whatcollectionsofhisremainareatthe R+&+ PhiladelphiaAcademyofNaturalScience.8,',(#(# Ї0  e Rolston,Holmes,III.(***).Contemporaryamateurbryologist,DistinguishedProfessorof  PhilosophyatColoradoStateUniversity, fatheroftheconceptofenvironmentalethicsas  amodernacademicdiscipline.CollectedbryophyteswithFredHermannforseveralyears,and | isanexcellentfieldbryologist.b(#(# 0  e Rothrock,JosiahTrimble(18391922).AstudentofAsaGray,andabotanistandsurgeononthe . ~ WheelerExpedition.HewasthesupervisorofJohnWolf.Manyvascularplantsbearhis  d name;whetherhereallycollectedbryophytesisquestionable. J(#(# 0  e _Sayre_,Geneva(19111992),CollectedmossesintheFrontRangeandinMesaVerdeNationalPark    andwrotebutdidnotpublishaMossFloraofColorado.Didextensivebibliographicwork    oncryptogamicexsiccati.See_Pfister_Ԁ(1993). (#(# 0  e Spence,Mrs.E.Jane.ReportedashavingcollectedatManitouandvicinityinJuly,1897(Ewan ^  1981).PossiblythecollectorofNeckeradouglasii?D (#(# 0  e _Stiverson_,ClareL.Bryologystudent,producedaMastersThesisfortheUniversityofDenverin ` 1951:ApreliminarysurveyofthemossesintheMountEvansarea,45pages.Someofher F collectionsareinherbariumCOLO.accessionnumbersB92639284.,(#(# 0  e _Wikel_,PatricianeeThomas(***).CollectedhepaticsintheFrontRangeandSanJuanMountainsin  the1970s.CollectionsareatCOLO.(#(# 0  e Wolf,John(18201877).Memberofthe1873Wheelerexpedition.AssistanttoRothrock.Several Z vascularplants(cf.Ribeswolfii)bearhisname.Itisquestionablewhetherhehadanyreal @ interestinbryophytes.&v(#(#   In19***theUniversityofColoradohostedaseriesoffieldtripsduringthefirst_ICSEB_ B meetinginBoulder,Alargenumberofbryologistsattended,includingWilliamC.Steere,Lewis ( Anderson,      _@: Habitatsofbryophytes   Wetlands: d @``>BogsandFens. 0    Coloradohasnotruebogs.WhathavebeencalledbogsinColoradoarereallyfens.McQueen  L (1990)describedthedifferences.Weparaphrasesomeofthesestatementstoapplytoour  2 mountainousregion.Bogsarewetlandsthatareverypoorinnutrientsbecausemostoftheminerals    areobtainedfromprecipitationratherthanfromgroundwater.Theyarealsoreferredtoas    ombrotrophicpeatlands,meaningsimplythatallnutrientscomestrictlyfromprecipitation.Theyare   usuallycharacterizedbyapHof4orless.Sphagnumisthedominantvegetationofabog. z    Fensareverysimilartobogs,buttheyreceivetheirnutrientsfromthesurroundingground F  waterandarewetter.Manyofthesameplantsthatarefoundinbogsarealsofoundinfens.Fensare ,|  oftenreferredtoasminerotrophicpeatlandsbecauseifthegreaterinfluxofnutrientsfromground b waterThetermfenmaybesomewhatmisleading,becausethecategoryincludessomediverse H habitats.Forexample,thewordfenisusedtodescribesomealkalinewetlandsthataredominatedby . sedgesandgrassesandveryfewpeatmosses.    Poorfensgenerallyobtainmorewaterfromprecipitationthanfromrunofffromelevated  places,andtheirpHishigher(5to6.5).HighCreekFen,inSouthPark,maybesocharacterized. v Richfensoccurinlevelareasintowhichwaterdrainsfromhigheraltitudesandflowsthroughand \ outalthoughataratherslowrate.ChattanoogaFen,intheSanJuanMountains,isanexample. B   Theextensivewillow/sedgemeadowsalongthesubalpinestreamsdonotholdwaterforlong ^ periodsandmaybecomequitedryseasonally.Therelativelymeagrenumberofbryophytesare D commonlyrestrictedtotheslightlyraisedbasesofwillowclumps. * ***Otherhabitatstocome,forexample)  Swampsareforestedwetlandsinwhichthereisstandingwayterformostoftheyear. r!  Carrsarewillowdominatedwetlandscommononmountainfloodplains.Thesearenotvery >#" productiveofbryophytesbecauseoferosivescouringbywaster,anddensityofCarexstands. $$t# Alkalineflats:seasonallywetgreasewoodstandswithfewspeciesofmosses,Crossidium,Syntrichia %@!% Gypsum/saltdomesthatretainwaterlongerthantheflatdesertpavements. ' #'   )$)    b @ 5 ThegeographyofColoradomosses     Mossesoftenhavemuchlargerormorewidelydisjunctdistributionpatternsthandoflowering ~ plants.TheSouthernRockyMountainsdrawsitsflorafromtheobviousmigrationpathwayafforded d bytheimmensenorthsouthextentoftheWesternAmericanCordillera.Everydrainagesystemthat J radiatesfromthecoreofthesystemalsoservesasahighwayofmigrationforriparianandlowland 0  ordesertspecies.Anaptanalogyisthatofagreatwheel,whosehub,theSouthernRockies,preserves  f themostancientsurvivors,andwhoseaxleandspokes"theCordilleraandthedrainagesystems,  L providethepathwaysalongwhichmigrationhastakenplacethroughtimeunderthepressuresof  2 climaticshiftsandorogenicmovements.      TheColoradomossfloraispredominantlyborealmontane,astestifiedtobythefactthatall   butaverysmallnumberofspeciesarecommontoColoradoandScandinavia. z    AsignificantnumberarerestrictedtoprotectedravinesintheeasternslopeoftheFront F  Range,wheretheypersistasrelictsofthePleistocene.Thesespeciesofeasternwoodlands(some ,|  havingextensionstothePacificNorthwest)formerlyradiatedtothefootoftheRockyMountains. b Followingthedryingoutofthehighplains,theywereeliminatedfromtheinterveningterritory. H AmongtheseareAmphidiummougeotii,Anomodonspp.,Brachytheciumacuminatum,Cnestrum . schistii,Dicranumbrevifolium,D.flagellare,D.fulvum,D.montanum,Didymodontectorum,  Entodoncladorrhizans,Grimmiaanomala,G.pilifera,Leskeapolycarpa,Mniumhornum,Neckera  complanata,Plagiotheciumcavifolium,P.laetum,Platygyriumrepens,Rhytidiadelphustriquetrus.    AfewdesertandlowlandspeciesarecommontodesertandsteppeareashereandinAsia \ (Anoectangiumhandelii,Didymodonanserinocapitatus,Jaffueliobryum,Crossidium,Aloina, B Pterygoneurum).Averysmallnumberofworldwidedisjunctscanbeconsideredtobeancientrelicts (x ofTertiaryorolderfloras(Bryoxiphiumnorvegicum,Gemmabryumalpinum,Leptodonsmithii,Oreas ^ martiana,Voitianivalis).AveryfewspeciesfollowaNorthAmericanCordilleraAndeanpath D rangingfromAlaskathroughColoradoanddowntosouthernSouthAmerica(Anacolia,Bartramia * potosica,Leptopterigynandrum,Rhexophyllumsubnigrum,Syntrichiabartramii).    Oftheborealmontaneelement,themajorityofourspeciesbelongtotaxaofrelativelydry   mountainareaswithcontinentalclimate.Speciesrequiringconstantlyhighhumidity(suboceanic r!  species)suchasthoseoftheforestsofthePacificNorthwestareeitherabsenthereortheyoccurin X"! smallremnantsofwhatmighthavebeenduringthePleistocenelargerareassuitedtotheirgrowth(, >#" Hylocomiastrumpyrenaicum,Hylocomiumsplendens,Pleuroziumschreberi,Ptiliumcrista $$t# castrensis,Rhytidiadelphustriquetrus).  %Z $ ***EndemicPacificNorthwest:Brachytheciumleibergii,Roelliaroellii... &&"& @C C ) Somelittlerecognizedfactsaboutbryophytedistribution (#(    Therearetwowonderfulthingsaboutbryophytedistributionswhichhavenothingmuchto r*%* dowiththebryophytesthemselves.ItisaboontoscientiststhattheLorddidnotdistribute X+&+ bryophytes,orplants,forthatmatter,otherplants,atrandom.IfHeorShedid,wescientistswould >,', haveaterribletime,becausewewouldhavetoscoureveryinchoftheplanet.Bryophytesaretied  verycloselytotheirmicrohabitats,whicharediscreteandcanberecognizedbyanastuteobserver.  Furthermore,bryologicalcollectorsmoveatasnailspacecomparedtowildflowerpeople.Wemight | walkafewmilesinaday"longerdistancesifwecloseoureyesuntilwereachthedestination b whereweknowwewillfindaparticularmicrohabitat.Usuallywetendtofindaninterestingspotat H theroadsideoronlyafewhundredyardsaway,andspendthedayrightthere! . ~   Acorollarytothefirstblessing,weshouldbeverythankfulthatmanhasbuiltroadsandtrails,  J despitethefactthatwetendtoresenttheseintrusionsonpristinehabitats.Butforthesewemight  0 neverfindsomeoftherarerspecies.Whenlookingatadotdistributionmapofabryophytewemust    rememberthatthisiswherecollectionshavebeenmade.Adotmapwillgiveusageneralideaof    otherlocalities,butfewareasaresowellcollectedthatonecanmakedetaileddotmapscoveringthe   smallergeographicaldivisionssuchascounties,townships,ranges,andsections.Thusitisnotvery x  meaningfultodeclareabryophyterarebecausefewcollectionshavebeenmade.Fewbryologists ^  collectthesamecommonmossinmorethanafewcountiesinastate.Atthesametime,becausewe D  havefoundararemicrohabitatdoesnotmeanthatwehaveexhaustedthepossibilitythatothersuch *z  placesmaystillbefoundfartherfromtheroad. ` @ + Somethoughtsaboutecology,evolution,andtaxonomy  ,   Bryophytesareobviouslycloselytiedtoparticularsubstrates,notonlythegeneraltypesof  rock,butthefeaturesofrocks.havingtodowithexposure,seepageareas,crevices;treeandshrub  habitats:thekindsofbark,theheightfromtheground,theexposure;soiltypesandconditions:soil v typesandconditions,whetherheavy,light,alkaline,compactedorloose;thesurroundingforest \ canopy.Othersmaycometomind. B   Koponen(1982b)presentsaninterestingdiscussionconcerninghowbryophyteecologymay ^ giveinsighttotheirtaxonomy.Hesays Itismucheasiertofindexamplesofspecieswhichare D strictlylimitedtoaspecialmicrohabitatthanofspecieswhichgrowinalargevarietyofhabitats, * suchasCeratodonpurpureus.Habitatspecificityofteniscorrelatedwiththeevolutionandhence  thetaxonomyofthespecies.Thisiswhyitissoimportanttoincludehabitatinformationwith  collections.WhenIwasidentifying, siteunseenDonaldMcVeansAustralianalpinecollections,   Isometimessuggestedaname. Itcantbethat;thehabitatisallwrong!Hewasrightahundred r!  percentofthetime.Koponensays. Iamalsoponderingwhyweshouldnotacceptaspecificstatus X"! forataxonwhichdiffersfromitsnearestrelativeinseveralcharactersandalsohasadifferent >#" ecology?Ifeelthatthisquestionispertinenttoourconsiderationofthetaxonomicpositionofthe $$t# highalpine varietyofAulacomniumpalustre.Thismossnevergrowswiththespeciesproper,its  %Z $ leafshapeisunique;itproducesnogemmae,itisdisjunctinhighmountainsoftheEastern %@!% hemisphere;itdoesnotfruit;itconcealswhatstemrhizoidsitpossesseswithintheaxilsoftheleaves; &&"& Ihavenotencountered intermediates. ' #' @: Geobotanyofbryophytes  n*%*   Becauseoftheirphysicalnature,theirexistenceaspredominantlyhaploidorganisms,andtheir <,', closeattachmenttotheverythinlayerofsubstrateonwhichtheygrowandmustsurvive,bryophytes,  morethananyotherplants,arerecognizedasbeingassociated,ifnotdependenton,variousphysical  substrates,oftenhavingtodowiththeavailabilityofwaterasactualliquidorasmistordew;if | liquid,whetherthewaterisrunningornot,whetheritisaugmentedbychemicalionscomingdown b fromahillsideorbyrunofffromminetailings,etc.,hatis,whatthepHofthesubstrateis,acidor H basicorneutral.Othercharacteristicsofthesubstratesonandinwhichtheygrowareimportant. . ~ Bryophytes,atleastmanyofthem,arecategorizedasbeingcalciphilesorbasiphiles(associatedwith  d calciumorothersubstancesincludingpotash,similarinchemicalbehavior);disturbanceplants,  J nitrophiles(requiringortoleratingnitrogenousmaterial);dung lovingmosses;minerotrophic  0 mosses.      Notverymanymossesarewellenoughknownforthemtobecategorizedwithanyofthese   terms,andmoreover,itisnotunderstoodwhattherolesofsubstratereallyplaysintheoccurrence x  ofmosses.Mossesthatareconsideredcalciphilesononepartoftheworldmaynotbehaveassuch ^  inanother.MossflorasinPlaceslikeScandinaviaaremorethoroughlyclassifiedthanothersbecause D  ofalonghistoryofinterestinthemandbecausecalciferousrockassociatedwithabundantmoisture, *z  makescalciferoussubstratesmoreobviousselectorsofplantspecies.IntheRockyMountains, ` however,wenotonlyhavehadlittleseriousinterestinmosses,andthesocalledcalciphilesareoften F notdefinitelyconnectedtocalciferoussubstrates.VerylittlenotehasbeenmadeinColorado , concerningthecalciumcontentofthesubstrate.    HereisapartiallistofColoradospeciesthatareconsideredbyCrum(1973)asbeing  calciphiles,orlikelycalciphiles.Afewmorehavebeenaddedbythepresentwriters.Thoseindicated t as(!Crum)weconsidersurprising,sinceinColoradotheydonotnecessarilybehaveassuch. Z e( X5(#(#(Abietinellaabietina(!Crum) &v Aloinarigida  \ Anoectangiumaestivum B Anoectangiumhandelii ( Anomodonrostratus  Anomodonattenuatus  Barbulaconvoluta   Barbulaunguiculata p!  Brachytheciumturgidum( probably!Crum) V"! Breidleriapratensis(!Crum) <#" Campyliumstellatum "$r# Catascopiumnigritum %X $ Conardiacompacta %>!% Cratoneuronfilicinum &$"& Cratoneuronfilicinum ' #' Dicranellavaria (#( Didymodonanserinocapitatus )$) Didymodonrigidulus l*%* Didymodonsubandreaeoides R+&+ Distichiuminclinatum 8,', Drepanocladusaduncus(!Crum) &v, Encalyptaprocera  \- Grimmiaanodon B. Gymnostomumaeruginosum (/ Homomalliumadnatum 0 Hygrohypnumluridum 1 Hymenostyliumrecurvirostrum  2 Hypnumcupressiforme(!Crum) p!3 Myurellajulacea V"4 Orthotrichumanomalum <#5 Orthotrichumpellucidum "$r6 Plagiopusoederiana %X 7 Pseudocalliergonturgescens %>!8 Pterygoneurumovatum &$"9 Rhytidiumrugosum(!Crum) ' #: Saelaniaglaucescens (#; Scorpidium(allspecies) )$< Sphagnumwarnstorfii l*%= Syntrichianorvegica(!Crum) R+&> Syntrichiaruralis(!Crum) 8,'? Tomentypnumnitens(!Crum)  Tortellatortuosa  Tortellafragilis | Tortula(Phascum) b Tortulaobtusifolia H Weissiacontroversa . ~   d   ( X5(@ 2Otherkindsofsubstraterelationships.  0 Disturbance:Ceratodon,Bryumargenteum,B.angustifolium,Gemmabryumsubapiculatum   Nitrophile:Ceratodonpurpureus x  Basiphile:Ditrichumflexicaule D  Minerotrophic:Coscinodoncribrosus,Mielichhoferia ` Fireash:Funariahygrometrica,Leptobryum , Dung:Splachnaceae(Tayloria,Splachnum,Voitia)  @} } / Somelittleobservedmorphologicalfeatures . t   Inanimportantpaper,Koponen(1982a)discussesmossrhizoids.Theirpositiononthestem B orleaves,andtheirsurfaceornamentation,haslargelybeenignored.Thecellsthataredestinedto (x producerhizoidsarealsolittlestudied,butinsomegroupslikeSarmentypnumandDrepanocladus ^ theyaretaxonomicallysignificant.Branchingpatternsofthestemalsovary.Thesepatternsareso D importantthatlateroninthisbookIshallmakeextensivequotesfromKoponenspaper. *    @ 3 KEYTOTHEGENERAOFCOLORADOMOSSES  (Inpreparation) @E  @ 3Family,Genus,andSpeciesTreatments  N N__Q_@> AMBLYSTEGIACEAE  6 Ref:Hedenspapers  Notethatseveralkeysfollowinsuccessionbeforethelistofgeneraandspecies.   @- - . FieldkeytothefenspeciesofAmblystegiaceae  f  @E  Forpeopleworkingwithfens,itisveryusefultobeabletoidentifytheplantsinthefield. j 4 Thiskeywillmakethejobeasier.Note:Leaffeaturesrefertothestemleaves,notthebranchleaves. P  1a.0  Leavesbroadlyovate,ovateorroundedtriangular,inupperpartsuddenlyroundednarrowed   orapiculate(note:theinflexedupperleafmarginsmaygivetheapexamoreamucronate   appearance)#@"@"x(#. #(#(##(2)   1b.0  Leavesstraightorfalcate,fromovateortriangularbasalportiongraduallynarrowedtowards   leafapex#@"@"x(#.r r $(#(##(5) ~  2a.0  Costashort,doubleorsingle,notorhardlyvisiblewithahandlens;shootsturgid,slightly J andirregularlybranched;alarcellsnothyaline,inanindistinctlydelimitedgroup. f0  Pseudocalliergonturgescens L(#(# 2b.0  Costalong,reachingleafmiddleorfurther,singleorbranched,usuallyeasilyseenwitha 4 handlens#@"@"x(#. !(#(##(3)  3a.0  Leavesusuallyovateornarrowlyovate;plantsofalpinepools.#@"@"x(#.W(#(##(4)  3b.0  Leavesusuallybroadlyovateorbroadlyroundedtriangular;greenorbrownishgreen  species,sometimeswithapalepinkishhue. Calliergon |(#(# 4a.0  Redcolorscommon,otherwisegreentodarkgreen;usuallystronglybranched;stemleaf J apexusuallydistinctlyapiculate,atleastinyoungleaves;leafpointoftenbentinwards;very f0 rarelyfoundwithleafbornerhizoids. Warnstorfiasarmentosa L (#(# _4b_.0  Paleoryellowgreen,usuallysparselybranched;leafapexroundedorroundedobtuse;leaf 4! bornerhizoidscommonneartheleaftips. Straminergonstramineum "(#(# 5a.0  Costashort,usuallydouble,notorhardlyvisiblewithahandlens#@"@"x(#.[(#(##(6) #! _5b_.0  Costalong,reachingmiddleofleaforfurther,normallysingle,usuallyeasilyseenwitha $" handlens#@"@"x(#. !(#(##(8) %~# 6a.0  Verylargespecies;shootsoftenturgid,notflattened;greenoroftenwithbrown,yellow 'J!% brown,red,orblackishcolors. Scorpidiumscorpioides f(0"&(#(# _6b.0  Mediumsizedspecies;shootsoftensomewhatflattened;greenorpaletoyellowgreen N)#' species#@"@"x(#.H H (#(##(7) 4*#( 7a.0  Leaveswithlargeanddistinctgroupsofalarcells;costashortandforked,orlacking.leaves ,%* stronglyfalcate Calliergonella(SeeHypnaceae) 6(#(# 7b.0  Leaveswithsmallandindistinctgroupsofalarcells;costaveryshortandinconspicuous;  leavesweaklyfalcate. Breidleriapratensis(SeeHypnaceae) (#(# 8a.0  Leavesstraight,lanceolate,stronglyplicate,theundersideofthestraightstemsdensely   clothedwithbrowntomentum. Tomentypnum  (#(# 8a.0  Notasabove#@"@"x(#.  $(#(##(9)  j 9a.0  Stemleavesusuallymoreorlessbroadlytriangulartoverybroadlycordate,ratherquickly l 6 narrowedtotheacumen;branchleavesmuchsmaller,falcate;alargroupslarge,triangular R  andwelldelimited,reachingfrommargintocosta;plantsusuallydenselypinnate;leavesnot 8  plicate. Cratoneuronfilicinum  (#(# 9b.0  Leavesroundedtriangular,ovateorbroadlyovatetolinear,moregraduallynarrowedtoward   theapex;alargroupsusuallylessdistinct;branchingrarelydenselypinnate(tomentummay   occurinPalustriellafalcata,Tomentypnum,andConardia).#!!w(#.R(#(##(10)   10a.0  Paraphylliapresent(tearoffafewleaves;theparaphylliaareusuallydistinctlyvisiblewith h ahandlens);largespecieswithfalcate,distinctlyplicateleaves;costastrong;tomentum N frequentlypresent. Palustriellafalcata j4(#(# 10b.0  Paraphylliaabsent;tomentumlackingexceptinTomentypnum#!!w(#.R(#(##(11) R 11a.0  Leavesconspicuouslyplicate;commonlyfruiting,thecapsulesnormallyhorizontal(not  commoninfensbutverycommoninwetspruceforests). Sanioniauncinata (#(# 11b.0  Leavesnotplicateorindistinctlyso#!!w(#.((=(#(##(12)  12a.0  Leavesfrommoreorlessstraightandrathererectbasalportionwithratherstronglycurved  upperpart;alargroupsundifferentiatedorverysmallandnotvisibleinthefield.#!!w(#. p(#(##(13) h 12b.0  Leavescurvedmoreorlessalongtheirentirelength,ornearlystraight;alargroupsdistinctly N differentiatedandusuallyvisibleontornoffleaves(indistinctlydifferentiatedin j4 Pseudocalliergonangustifolium)#!!w(#.LL7(#(##(14) P  13a.0  Shootsrelativelystiffandmoreregularlybranched;green,yellowgreen,brown,orbrown " red(auniquecolorcombinationincontrasttothenext);leavesgreenoryellowgreenwith #  brownredcosta(oftenalsoleafbase),somewhatdull(duetorelativelyshortleafcellswith #! squarishends). Scorpidiumcossonii $"(#(# 13b.0  Shootssomewhatlargerandlessbranched;red,blackishred(brownred)orgreen;glossy %# (duetolongcellswithgraduallynarrowedends. Scorpidiumrevolvens &f $(#(# 14a.0  Costalongexcurrent;shootandbranchapicessometimespencillike;usually_growing j(4"& submergedinpools#!!w(#.& & *(#(##(15) P)#' 14b.0  Costanotexcurrent"incaseofdoubt(occurringinspecieswithradiallybranchedshoots) 6*$( theshootandbranchapicesarenotpencillike;submergedornot#!!w(#.ddY(#(##(16) +$)  ,%* 15a.0  Shootandbranchapicessometimespencillike;leavesstraightorcurved,oftendeepred 6 purple;youngaxillaryhairslongandbrown(visiblewithahandlensifafewleavesaretorn  off);shootsradiallybranched. Warnstorfiatrichophylla (#(# 15b.0  Shootandbranchapicesneverpencillike;leavesusuallycurved,neverbecomingred;young  axillaryhairssmall,hyaline(notvisiblewithahandlens);shootsdistichouslybranched.    Drepanocladuslongifolius  (#(# 16a.0  Yellowbrown,brownishyellow,orgreenspecies,typicallywithagoldenglosswhendry;  N alarcellsindistinctlydifferentiated;plantsofstronglycalcareoushabitats. Pseudocalliergon j 4 angustifolium R (#(# 16b.0  Colorvarying,notwithgoldenglosswhendry;alargroupsusuallylargeandmoreorless :  distinctlydifferentiated;inlessstronglycalcareoussites#!!w(#.T(#(##(17)    17a.0  Shootsdistichouslybranched,neverred;rhizoidsnevergrowingfromtheleaves;mostlyin   nutrientrichhabitats. Drepanocladus  (#(# 17b.0  Shootsmoreorlessradiallybranched,mostdistinctwhengrowingwithsteminvertical   position;sometimeswithredcoloration,andsometimeswithrhizoidsgrowingfromthe j leaves;inlessnutrientrichsituations. Warnstorfia P(#(# 18a.0  Planthabit Drepanocladuslike,thatis,withleavesfalcatesecundtosometimesstraight T anderect;leafacuminainstraightleavedplantsplaneoratmostslightlyfurrowed. :  Drepanocladusaduncus  (#(# 18b.0  Planthabit Campyliumlike,thatis,withleavesfromstraightanderectbasesusuallywith  leafacuminamoreorlessspreadingorsquarrose;leafacumenfurrowed. Drepanocladus  polygamus (#(#  @' ' . Hintstorecognitionofgeneraofflowingwater  T   Aquaticpleurocarpousmossesthatgrowoverseepingrocksorattachedtorocksinsmall X " streamsmaybelongtogeneraofseveralfamilies.Brachytheciumrivularemaybemistakenfor >! Amblystegium,buttheleavesaremorebroadlyovateandfinelyserrulatedistally,andthealargroup $" isconspicuouslyinflated.Amblystegiumripariumisnotoriouslyvariableinthesize,lengthand  #  shapeofitsleafcells,butthemarginisentire,thealarcellslesspronounced,anddoesnothavethe #! rather_stiffish_Ԁtextureofthebranches.Nevertheless,Ihavetroublewiththese. $"   ItseemstobeaHobsonschoiceofrecognizingthesmallergenerainthelargegenus &l $ Amblystegium.ItendincreasinglytoagreewithCrumandAndersonwho reluctantlydecidedto 'R!% treatHygroamblystegiumandLeptodictyumaspartofAmblystegium.Althoughtherearegreat n(8"& differencesinsizeandappearancebetweenthetineA.serpens(whichgrowsatthebasesoftreesand T)#' shrubs),theBrachytheciumlikeA.riparium,theharshlystiffHygroamblystegiumofrunningwater, :*$( thereseemstobenoreallyqualitativecharactersetstoseparatetheseasgenera.  +$)  ,%*   Hygrohypnumisrathereasytorecognizeasagenusalthoughitsspeciesarereallydifficult 6 todistinguish.  @@ Generalkey   la.0  Alarcellsinflated,sharplydifferentiated;paraphylliausuallypresent;leafcellsmoreorless   papillose(fromprotrudingcellends). Cratoneuron and Palustriella  f(#(# lb.0  Alarcellsvarious;paraphylliaabsent;leafcellssmoothornearlyso#!!w(#.88^(#(##(2))  N 2a.0  Leavesusuallysquarrose. Campylium andrelatives (SeeCampyliaceae) P (#(# 2b.0  Leavesneversquarrose#@"@"x(#.VV.(#(##(3) 8  3a.0  Leaveswithasinglecostaextendingatleasttomidleaf#@"@"x(#.Q(#(##(4)   3b.0  Leaveslackingacosta,orthecostashortanddouble#@"@"x(#.rrM(#(##(8)   4a.0  Leavescordateovate,oblongovate,oroblong,neverlongacuminate;marginsentire.    Calliergon anditsrelativesf(#(# _4b_.0  Leavesalwaysdistinctlyacuminate;marginseitherentireorserrulate#@"@"x(#.^(#(##(5) N 5a.0  Leavesusuallyfalcate(exceptinsomesubmergedforms)orplicate,orboth. Drepanocladus P  anditsrelatives8(#(# _5b_.0  Leavesneitherdistinctlyfalcatenorplicate#@"@"x(#.zzE(#(##(6)  6a.0  Costastrong,percurrentorexcurrent;plantssubmergedinwater. Amblystegium   ( Hygroamblystegium)(#(# 6b.0  Costashorter;plantsterrestrialorinwetplaces,rarelysubmerged#@"@"x(#.\(#(##(7)  7a.0  Leavessmall(1!1.2mmlong);alarcellsrectangular. Amblystegium ( Leptodictyum)N(#(# 7b.0  Leaveslarger,or,ifsmall,thealarcellsotherwise. Amblystegium l6(#(# 8a.0  Leavesandbranchesverysmallandslender,leavesupto0.45mmlong,ovatelanceolate, :! neversquarrose,entireoroftenserrulateatthebase;2-3celledgemmaepresentintheleaf  " axils. Platydictya (seeHypnaceae)# (#(# 8b.0  Leaveslarger,orifsmall,thensquarrose#@"@"x(#.B(#(##(9) #! 9a.0  Leavesovatetoovatelanceolate,acuminate,usuallysquarrosebutsometimesmerelywide %# spreading. Campylium anditsrelatives ( See Campyliaceae) &j $(#(# 9b.0  Leavesotherwise,nottaperingtoanarrowapex#!!w(#.G(#(##(10) 'R!% 10a.0  Leavessmallorlarge,lessthantwiceaslongaswideor,iflonger,thensomewhatfalcate; T)#' basalcellsrarelywithpittedwalls;onwetrocksinornearstreams. Hygrohypnum :*$((#(# 10b.0  Leavesusually2mmlongorlonger,commonlytwiceaslongaswideormore,deeply "+$) concave,neverfalcate;wallsofbasalcellsoftenpitted;notattachedtorocks,buteither ,%* submergedinfenpondsorinloosegravelofsnowmeltrills0 (#(#  0  (# (#  #!!w(#.Z (# (##(11)  6  11a.0  Leavesmoreorlessfalcatesecund,atleastthoseofthetipsofthestemsandbranches;alar  cellshyaline,thinwalledandinflated,insmall,inconspicuousgroups. Scorpidium (#(# 11b.0  Leaveslooselyimbricatetospreading,notatalloratmostveryslightlysecund;alarcells   shortlyoblongand_subquadrate_,neitherhyaline,thinwalled,norinflated. Pseudocalliergon  (#(# @B B 0 Amblystegium Bruch&Schimper,1853 ,proper  P 1a.0  Costalessthan3/4theleaflength. A.serpens T (#(# 1b.0  Costaextendingintotheacumen,sometimesfillingitorexcurrent. A.varium < (#(# @ 4 Amblystegium ( Hygroamblystegium)    1a.0  Plantslarge,coarse,usuallyrigid;costaverybroad,110140mwide,terete,fillingtheleaf   tipandexcurrentasashort,stoutpoint. A.noterophilum  (#(# 1b.0  Plantssmallormediumsized,usuallyrelativelysoft;costalessthan100mwide,usually p somewhatflattened,endingintheleaftiporexcurrentasaslenderpoint#@"@"x(#.b(#(##(2) V 2a.0  Leavesbluntpointed;costarelativelybroad,75100mwide;upperleafcells36:1;cells X" attheinsertiongreenish. A.fluviatile >(#(# 2b.0  Leafapexacuteoracuminate;costalessthan75mwide;basalcellsmoreorlessyellowish. &  A.tenax (#(#  @7 Amblystegium ( Leptodictyum)  la.0  Plantscommonlysubmergedinwater;leavesentire,usuallyattachedobliquelytothestem, t thebranchessomewhatcomplanate;medianleafcells50l00mx5l0m(relativelylong Z andslender). A.fluviatile v@(#(# 1b.0  Plantsnotusuallysubmergedbutofteninverywetplaces;leavesentireorserrulate;leafbase ^ ( attachedtothestemtransversely(bowedoutatthebase),theupperpartoftheleaferect D! spreadingandsomewhattwisted;leafcells30-60mlongx5mwide(relativelyshort). A. *" riparium # (#(#   A.fluviatile (Hedwig)Bruch&SchimperThusfarwehaveonlyonerecordofthis,fromthe $" LittleRoyalGorgeofNorthBoulderCreek. %#   A.riparium (Hedwig)Bruch&Schimper.Commoninverywetplaces,particularlyinareas '^!% wherelivestockorotherpastoralactivityisstronglyevident.Ihavefounditabundantindrinking |(F"& troughs,oncheckdamsluicesandinrelativelystillwaterthroughthemiddlealtitudes. b),#'   Inourdistributionofspecimens,wemadeahorrendousmistakeofthinkingalarge,robust, .+$) submergedpopulationwasFontinalishypnoides.Evidentlywewerenotalone.Crum&Anderson, ,%* p.994,write: Amblystegium_laxirete_representsanextremedevelopmentofaFontinalislikehabit. 6 Itsometimesoccursinlong,streamingmassesinswiftlyflowingwaters....Inthishabitat,theplans  arestriking,butinlessvigorouslyflowingwaterstheyarelessdistinctive.Anumberofother  namedformsevidentlyarevariantsofthisextraordinarymodification.Moral:Dontassumethat  everythingthatlookslikeFontinalis,isFontinalis!     A.serpens (Hedwig)var. juratzkanum (Schimper)Rau&_Hervey_.Asmall,nondescript  d species,usuallyfruiting.Theleavesareabout0.6mmlongandgracefullyacuminate,withcostato  L aboutmidleaf.Themedianleafcellsareabout6:1or35mx5-7m,smoothandmoderatelythick h 2 walled.Theleafmarginisslightlydenticulatefromprojectingdistalendsofthecells.Thealar N  regionisquiteclearlydifferentiated,consistingofquadrateandbroadercellsformingamoreorless 4  triangularpatchatthebasalangles.Thecapsuleiscurved,theoperculumconic,andtheurnstrongly   constrictedbelowthemouth.Thespeciesisgenerallydistributedonwetboulders,soil,basesof   saplingsorshrubs,exposedroots,rottenlogs,etc.,fromthefoothillsuptothesubalpine.Itcanbe   mistakenforsmallspeciesofCampylium,especiallyCampylophyllum.hispidulum.Theleavesare   neithersecundnorfalcate,andgenerallydonotspreadverywidelyfromthestem,slightlymore |  whenwet.ConardiacompactamayalsobetakenforAmblystegiumserpens.Iaminclinedtofollow b NyholmintreatingA.juratzkanumasavarietyofA.serpens.Thevarietyisdistinguishedbyhaving ~H theleavesmorewidelyspreadingandwiththemarginalbasalcellsrectangularinsteadofquadrate. d.   ***deleteA.tenax (Hedwig)C.JensenAfrequentlyencounteredspeciestightlyattached 0 torocksinthelowerfoothillcanyons.AsfarasIknow,itsaltitudinalrangeisquitelimitedin  Coloradototheareasnearthemouthsofthefoothillcanyons.Itisacoarse,harshmosswithwiry  stems,andtheolderpartsaredenudedofleavesasinthelargerHygrohypnumspecies.Thecostais  extremelystout,however,andpersistsasaspinelongafterthecellsofthelaminahavebeenworn  away.InHygrohypnumthecostaisneversowelldevelopedandresistanttowatererosion.The z stemsandleavesaccumulatebitsofsandandcrustaceanshells,diatoms,etc.,andoftenhaveavery ` grittyfeel. |F   A.varium (Hedwig)LindbergWehaveonecollectionthusfar,ofthisspecies,whichshould H  befairlycommoninloweraltitudes:MontroseCo.:_Nyswonger_ԀMesa,aboveandwestofParadox, 0! ca1mibelowBuckeyeReservoir,inpineforest,7,200ft.alt.,N38$24-47.W109$02-19.;atedge " ofverysmallstreamletatthebaseofrimrock,2July2000,Weber,Wittmann,&LehrB112608. "  Itisasmallspecies,likeA.serpens,butthecostaisverystoutandfillstheacumen. #! @  - Calliergon (SullivantinA.Gray)Kindberg,1894 %x#   Calliergonisgenerallynotaplantoftheedgesofswiftlyflowingbrooks.C.richardsonii |'F!% growssubmergedinstillpools.C.cordifoliumoccursonsaturated,swampygroundinforest b(,"& clearingsfilledwithtallwillows,C.giganteumwasfoundonslopingrockfacesbesideaquiet H)#' backwateronalevelbenchofamountainstream.AfewspeciesformerlyincludedinCalliergonare .*#( commoninalpinepools.TheseincludeWarnstorfiasarmentosaandStraminergonstramineum. +$)  +%* 1a.0  Costausuallyendingwellbelowtheleafapex,withshortbranchesorforkedattheapex; 6 shootswithratherlong,thickbranches,moresparselybranchedandwithbranchleavesmore  erectorimbricatethaninC.giganteum(seebelow). C.richardsonii (#(# 1b.0  Costaendingalmostintheleafapex(appearingtoreachtheapexasseenwithahandlens)  #@"@"x(#.(#(##(2)   2a.0  Costastrong;alargroupsofstemleaveslarge,triangularandsharplydelimitedfrom  f surroundingcells,extendingfromleafmargintooralmosttothecosta;leavesbroadly  L triangular(shootswhenwelldevelopeddenselybranchedlikeasprucetree,withmoreor h 2 lessspreadingbranchleavesexceptnearbranchapices. C.giganteum N (#(# 2b.0  Costaweaker;alargroupssimilarbutdiffuselylimitedfromsurroundingcells;rarelyfound 6  inpermanentlysubmergedhabitats). C.cordifolium  (#(#   C.cordifolium (Hedwig)Kindberg.Poolsandlakesideswamps,uppermontaneand   subalpine.OurrecordsarefromtheplateausofwesternColoradoandfensintheFrontRange.     C.giganteum (Schimper)Kindberg.Wehavetwocollections:LarimerCo.:Cirque h Meadowstrail,vic._Pingree_ԀPark,streamlettributaryofFallCreek,9,600ft.,Hermann&Rolston P 80114.Samelocality,Weber&Wittmann112731. l6   C.richardsonii (Mitten)Kindberg.Submergedinfens,Boulder,Larimer,andClearCreek 8 counties.TheBoulderCountyspecimensareextremelylargeleaved,resemblingC._megalophyllum_   (soreportedbyWeber,1973).  @ 3 Cratoneuron (Sullivant)Spruce,1867    C.filicinum (Hedwig)Spruce.Thisspeciesisabundantinwetsitesinthesubalpineforests j andwillowcarrsandfens.Theplantsaresmall,pinnatelybranched,andthestemleavesare R conspicuouslylargerthanthebranchleaves.Thebranchleavesareusuallyfalcate.Acharacteristic n8 featurearetheparaphyllia,ofvariousshapesandsizes,minutely_leaflike_,whichunfortunatelyvary T  inabundanceandaresometimesalmostlacking.Thestemleavesarebroadlyovate,_witrh_Ԁavery :! strongcosta.Themedianlaminalcellsarenarrowlyrhomboid,anthealarcellsareenlarged,ina  " conspicuoustriangulargroup.Thespeciesissaidtobestrongly_calciphilous_,andiscommonin #  calcareousfensinthesubalpine. #! @d d 2 Drepanocladus (MllerHal.)Roth,1899 %#   CrumandAndersonacceptDrepanocladusandScorpidiuminatraditionalsense,whichfor 'P!% fieldbotanistsseemsreasonable,andtheyrejecttheseparationofseveralotherspeciesintothe l(6"& generaLimprichtia,Warnstorfia,andScorpidium.However,wefollowtherecentworksofHedens. R)#'   HedenssuggestsonewaytodistinguishDrepanocladusfromWarnstorfia: Itisalways +$) goodtolookforrhizoidalinitialsneartheleafapices.Ifyoustudy1015leaves(orashootapexwith ,%* severalleavesleft)inthemicroscope,therearealmostalwaysatleastsomeleaveswithsuchinitials 6 inWarnstorfiaspecies,butneverinDrepanocladuss.str.  1a.0  Leavesfrommoreorlessstraightanderectbases,usuallywiththeacumenmoreorless  spreadingorsquarrose;leafacumenfurrowed;plantsmoregoldenbrownthangreen. D.   polygamus  (#(# 1b.0  Leavesstronglyfalcatesecund;leafacumeninlessfalcateleavedplantsplaneoralmost  h slightlyfurrowed;plantsgreenwithnoothertints#@"@"x(#.K(#(##(2)  N 2a.0  Costaofstemleavesendingwellbelowtheleafapex;leafmarginentire,oronly P  occasionallyveryfinelydenticulate. D.aduncus 6 (#(# 2b.0  Costaofstemleavesexcurrent,rarelyendingafewcellsbelowleafapex;oneorbothleaf   marginsusuallypartlyfinelydenticulate. D.longifolius  (#(#    Drepanocladusaduncus (Hedwig)Warnstorf.Anextremelycommonandvariablespecies,   verywideranginginaltitudefromtheplainsuptothealpine.Theelongate,inflatedalarcellsina   rowcontinuingalmosttothecosta,isdiagnostic.Theplantisweak,notstandingerect,yellowish j green,neverwithreddishorbrowncolors,verylittlebranchedandthennotatallpinnate;thecosta P isslenderandelongate.However,somecollectorshaveconfusedthiswithCalliergonella(was l6 Hypnum)lindbergii.Thelatterstandsstifflyerectincloseorder,theleavesareverybroadatthebase R (triangularovate)andwithlargealarcellsatthebasalanglesjustwheretheleafbecomesdecurrent. 8 Thecostaisabsentorshortanddouble,andfaint,andtheleafmarginstendtocurveinwardnearthe  apex.    Drepanocladuslongifolius (Mitten)Paris.Insubalpinepools.Wehaveoneexcellent  collection:GrandCo.:RockyMt.Nat.Park:FloatingonpondalongUteTrailsouthofLakeIrene,  10,400ft.,Hermann26506.Hedens(_BFNA_)acceptsthisforColorado,saying: D.longifolius[D. h _capillifolius_ofauthors]differsfromallotherAmericanDrepanocladusspeciesinitsexcurrentleaf N costa.BecauseofthelatteritcouldhardlybeconfusedwithanyearlierDrepanocladuss.l.inNorth j4 AmericaexceptWarnstorfiatrichophylla.However,theleavesarealwaysgreen,thereareneverany P  rhizoidalinitials,andtheshootsaredistichouslypinnatelybranched.InWarnstorfiatrichophyllathe 6! shootsareradiallybranched,andtheshootsandbranchapicespencillike,theleafmarginsaremore " stronglydenticulatethaninD.longifolius;itfrequentlybecomesredwhenemergentwhereasD. #  longifoliusnevergetsred,andtheaxillaryhairsconsistof17earlybrownuppercells(12elongate #! hyalinecellsinD.longifolius). $"   Drepanocladuspolygamus (Bruch&Schimper)Hedens.Onerecord,fromasubalpinefen &d $ onDiamondLake,intheFrontRange,collectedbyHermann(!Lawton).SeeFlowers,plate119:9 'L!% 11.(Campyliadelphus,Campylium,Drepanocladus).Thishasuntilrecentlybeenplacedin h(2"& Campyliumandindeedhasmoretheaspectofthatgenus.Theplantisdefinitelystouterandthe N)#' leaveslessfalcatethanD.aduncusandhasmorebrownishtints.D.polygamousisautoicous,while 4*#( D.aduncusisdioicousbutthisisdifficulttodeterminewithoutcapsules.