Promotor: dr.J.J .Barkman ,hoogleraa r ind evegetatiekund ee nbotanisch e oecologie t\)Ti Gio\ <9sig Ö
AnnaElisabet hJanse n
Thevegetatio nan dmacrofung io faci doakwood si nth eNorth-Eas tNetherland s
Proefschrift terverkrijgin gva nd egraa dva n doctori nd elandbouwwetenschappe n opgeza gva nd erecto rmagnificus , dr.C.C .Oosterlee , hoogleraari nd eveeteeltwetenschap , inhe topenbaa rt everdedige no p vrijdag2 oktobe r198 1 desnamiddag so mvie ruu ri nd eaul a vand eLandbouwhogeschoo l teWageninge n
• /ƒ.'/• Nb ClCj-oi BIBLIOTHEEK LH. 2 9 SEP. 1981 ONTV. TIJDSCHR. ADM M.{-*> w '?<~.s^ Stellingen 1. De ruimtelijke frequentie van carpophoren is een betere maat voor de abundantie van fungi dan de abundantie van carpophoren. 2. Om te voorkomen dat veel fungi die typisch zijn voor het Dicrano-Quercetum daar zullen verdwijnen als de strooisel- en humuslaag te dik wordt, zal strooiselroof moeten worden toegepast. 3. In bossen op voedselarme zandgrond is de dikte van de humuslaag meer bepalend voor de voedselrijkdom dan de concentraties van de ionen in de humus. 4. Het Luzulo-Fagetum leucobryetosum en het Fago-Quercetum dicranetosum dienen, net als het Dicrano-Quercetum, in het Dicrano-Pinion te worden geplaatst. Lisiewska.M.(1974): Macromycetes of beech forests within the eastern part of the Fagus area in Europe. Acta Mycol. 10: 3-72. Jahn.H., A.Nespiak & R.Tüxen (1967): Pilzsoziologische Untersuchungen in Buchenwäldern (Carici-Fagetum, Melico- Fagetum und Luzulo-Fagetum) des Wesergebirges. Mitt. Flor.- soz. Arbeitsgem. N.F. 11/12: 159-197. 5. De keuze van Agaricus dryophila Fr. als typesoort van het genus Collybia is een ongelukkige keuze, daar dit tot gevolg zal hebben dat Marasmius sectie Androsacei (M. androsaceus en M. splachnoides) onder Collybia gerekend moeten worden. Lennox,J.W. (1979): Collybioid genera in the Pacific Northwest. Mycotaxon 9: 117-231. Clémençon,H. (1981): Kompendium der Blätterpilze I. Collybia. Z. Hykol. 47: 5-25. 6. Aangezien biotoopverstoring, b.v. door bemesting of ontwatering, en het verdwijnen van groeiplaatsen, b.v. door bebouwing, meer bijdragen aan de achteruitgang van de paddestoelenflora dan plukken (waarvan de schade uitsluitend visueel is) zijn plukverboden niet zinvol en zeker niet afdoende om een verdere achteruitgang van de paddestoelenflora te voorkomen. Hethooghoude n vand ereeënstan ddoo r bijvoedering ind ewinte r lijktmee ro pextensiev e veehouderij dano p natuurbehoud. Het ideeda t aquatischemilieu s altijd "eenvoudiger" zijnda n terrestrische is,zeke rwaa r heteutroof , hydrologisch nietafgeslote nwate r inNederlan dbetreft , eenhardnekki g vooroordeel. Hetgebrui k demenselijk e bevolking uitt edrukke n in eenaanta l "zielen"dien tvermede n teworde ndaa r dit uitgaatva n een bepaalde,zee rbeperkt e interpretatie vanhe t begrip ziel endaaro m kwetsend kan zijnvoo r personend-i ehierove r een andere opvatting hebben. Stellingen bijhe tproefschrif t van A.E.Jansen, Thevegetatio n andmacrofung i of acid oakwoods inth e North-East Netherlands. Wageningen,2 oktobe r 1981. AnnaElisabet hJansen .Agricultura lUniversity ,Wageningen ,sectio nBiolog icalStation ,Kampswe g 27,941 8PD ,Wijste r (Netherlands).Th evegetatio n andmacrofung io faci doakwood si nth eNorth-Eas tNetherlands .Thesis .1981 . 131pp ,1 1Tbs ,4 0Figs .Engl. ,Ned .inl .e nsamenv . The fungusvegetation ,th evegetatio no fphanerogams ,mosses ,an dlichens , the soilprofiles are described and soil chemicalanalyse sar emad ei nth e associations Dicrano-Quercetum,Querco-Betuletum ,Violo-Quercetum ,an dit s sub-associationilicetosu m (sub-ass.nov. )i nth eprovinc eo fDrenth e(North - EastNetherlands) .Ne wmethod s areapplie d inth e soil chemical analysis (amounto f ionsexpresse d inmas s concentration,calculatio no ftota lio n supply), inth evegetatio nanalysi s (separateanalysi so fth emos ssynusi a onrotte n treestump san ddea dwood) ,an di nth emycosociologica lanalysi s (determinationo fspatia l frequency).Th elast-name dmetho di suse dfo rth e determinationo restimatio no fth eabundanc eo fth emyceli ape rspecies ,o f theplo thomogeneity ,o fth epatter no fth especies ,an do fth ecorrelatio n betweenspecies .Base do nth ephytocoenos eth eQuerco-Betuletu m andth eVi olo-Quercetum appearedt ob eclosel yrelated ,an dt obelon gt oth eallianc e Quercion robori-petraeae. Itwa s confirmed thatth eDicrano-Quercetu m is lessrelated , and did notbelon gt othi s alliance. Inth esoi lther ear e differences between the syntaxai nth eprofile san di nth e 'totalio nsup ply'. Th e syntaxa haveman y differential speciesamon gth efung ii na wa y to support the concepts onth e affinitiesbase do nth ephytocoenoses .Th e 'minimal area'o fth e fungusvegetatio nwa sdetermined .Distributio npat terns of fungus specieswithi nplot sar ediscussed .Annotation so nidenti - faction and taxonomy of fungi aregiven ;a ne wfungu staxo ni sdescribed : PsathyrelJa fulvescens var. dicrani. VOORWOORD Ditonderzoe ki sal see npromotie-onderzoe k aand eLandbouwhogeschoo lt e Wageningen,vakgroe pVegetatiekunde ,Plantenoecologi ee nOnkruidkunde ,sec tieBiologisc hStation ,uitgevoer di nd eperiod eaugustu s197 6- apri l1980 . Debegeleidin gberustt ebi jprof.dr .J.J .Barkman . Voorhu ntoestemmin g onderzoekt emoge ndoe ni nbosse ndi ehu neigendo m ofi nhu nbehee rzijn ,be ni kdan kverschuldig d aand eVerengin gto tbehou d vanNatuurmonumenten ,he tStaatsbosbehee ri nd eprovincie sDrenth ee nOver ijssel,mevrou wMoret , deeigenaar sva nhe tAmshoffsbos ,d eopzichte rva n het landgoed "DeEese" ,d egemeente sAsse ne nHoogeveen .Mij nhartelijk e dank gaat uit naar devel epersone n diemi jdaadwerkelijk e hulpgaven . Zonder diehul p zou ditproefschrif t nooitzij n gewordenwa the tn uis . Dr.E .Kit s van Waveren, dr.C .Bas , F.Benjaminsen , T.Boekhout , T.W. Kuyper, dr.R.A .Maa s Geesteranus, dr.M.E .Noordeloos , J.Schreurs ,me vrouw G.D.Tjallingii-Beuker s determineerden velepaddestoele nvoo rmij ; N.Deodatu s deed alle chemische analysesva n debodem ,H .Klee smaakt ed e tekeningen, mevrouw S.Kerkhove n corrigeerde deengels e tekste nverde r dank ik B.va nHulzen , E.Melis , J.va nReissen . Drs.P .Ypelaa r endr . J.J.Barkma n dank ikvoo rhu ntoestemmin g gebruik temoge nmake nva nno g niet gepubliceerde gegevens.Bijzonder e dankbe n ikverschuldig d aanmij n vader,Piete rB .Janse ne naa nHenri kW .d eNie ,di eel ko phu nmanie rmee r dedenda nallee nhe tdeterminere nva npaddestoele ne nhe tgeve nva nstatis tisch advies enhulp .Mij ncollega' sva nhe tBiologisc h station danki k voor hunbetrokkenhei d inplannin ge nuitvoerin gva ndi tonderzoek .I nhe t bijzonder mijnpromotor ,prof .dr .J.J .Barkman , diehe tmogelij kgemaak t heeftda t ik ditonderzoe k kon gaandoe ne ndi eme tonvermoeibaa renthou siasme, interesse enwerklus t ditonderzoe k begeleidheeft .Voora lj ebe trokkenheid inhe t afgelopenjaa rbi jd efas eva nuitwerke ne nopschrijve n zal ikmi j alszee r stimulerend blijven herinneren.Tenslott e danki kd e afd.Tekstverwekin gva nd eL Hvoo rhe tverzorge nva nhe ttypewerk . CONTENTS Inleiding 1 Samenvatting •5 1 Introduction 8 2 The Phyt ocoenoses 11 2.1 Introductionan dmethod s 11 2.2 Results 21 2.2.1 TheQuerco-Betuletu m 21 2.2.2 TheViolo-Quercetu m 23 2.2.3 TheViolo-Quercetu milicetosu m 25 2.2.4 TheDicrano-Quercetu m 26 2.2.5 Classificationi nhighe rrank s 27 2.2.6 Mathematicalaffinitie s 28 2.3 Conclusions 30 3 The Soil s 31 3.1.1 Introduction 31 3.1.2 Generaldescriptio no fth esoi li nDrenth e 31 3.2 Soilprofil s 32 3.2.1 Methods 32 3.2.2 Results 35 3.3 Soilchemica lfactor s 37 3.3.1 Methods 37 3.3.2 Results 38 3.3.2.1 pH- freshan dpH-H 20 38 3.3.2.2 pH-KCl 38 3.3.2.3 Conductivity and 'total'io nsuppl y 40 3.3.2.4 Specificgravit y 42 3.3.2.5 Na,K ,Mg ,an dC aion s 43 3.3.2.6 Totalcarbo nan dnitroge n 47 3.4 Conclusions 47 4 The Mycocoenoses 48 4.1 Introduction 48 4.2 Methods 48 4.2.1 Relevésan dtable s 48 4.2.2 Theincreas ei nth enumbe ro fspecie spe r yearo finvestigatio n 51 4.3 Results 55 4.3.1 Numbero fspecie s 55 4.3.2 Differential species 55 4.3.2.1 Introduction 55 4.3.2.2 TheDicrano-Quercetu m 56 4.3.2.3 TheQuercio nrobori-petraea e 58 4.3.3 Mathematicalaffinitie s 63 4.3.4 Representativeplo tare a 63 4.4 Conclusions 66 5 Annual Frequency 67 6 Spatial Frequency 68 6.1 Introduction 68 6.2 Methods 68 6.3 Results 69 6.3.1 Introduction 69 6.3.2 Frequencyhistogram san dhomogeneit y 69 6.3.3 Therelatio nbetwee nth efrequenc yan dth e abundance 73 7 Distribution patterns 76 7.1 Introductionan dmethod s 76 7.2 Distributionpattern s 76 7.3 Correlateddistribution s 81 8 Composition of ecological groups 92 Summary • 95 Literature 98 Appendix 1 Listo fplot s 103 2 Annotationso nidentificatio nan dtaxonom y ofsom efung i 107 Index on fungi 124 INLEIDING Vanuithe tBiologisc h Stationt eWijste r (Drenthe)word t alzee rvel e jarenee nonderzoe kgedaa nnaa rd esociologi ee nd eoecologi eva npaddestoe len.Barkma ne nD eVrie sdoe ndergelij konderzoe kaa njeneverbe sstruwelen , Arnolds aan graslandvegetaties ,e nenkel eander emedewerker se nstudente n aan sparrebossen, droge heidevelden, en eikenkreupelbosjeso pvoormali g stuifzand. Het uiteindelijke doel isd epaddestoelenvegetatie s vanall e vegetatietypeni nhe tDrent sdistric tt ebeschrijven ,zowe lkwalitatie fal s kwantitatief. Daarmee hopen we eenvolledi g inzicht ind e sociologische amplitudo en statuse ni n deoecologisch e amplitudova n depaddestoelen soorten inDrenth e teverkrijgen .Oo kverwachte nw e datdez ekenni sza l bijdragento the toplosse nva nvel esynsystematisch eproblemen ,speciaa li n dievegetatie sme tmaa rweini ghoger eplante ne nmossen . Onderzoeknaa rpaddestoelenvegetaties ,meesta lmycosociologisc ho fmyco - coenologisch onderzoek genoemd,word t inNederlan d alleenbedreve nvanui t hetBiologisc hStatio nt eWi jster .Buite nNederlan dvind tversprei di nWes t Europa enig onderzoek plaats. Onderzoek dato p eensystematisch emanie r allevegetatie-type ni n eenbepaald e streek omvat,vind teigenlij k alleen plaatsi nOos tEurop a (Polen,Hongarije ,Tsjechoslowakije) . Overhe t algemeen isno gmaa rweini gbeken dva npaddestoelenvegetatie s inassociatie s vanhoger eplanten .D emycosociologi ei see nno gvri jjong e tak van wetenschap die tamelijk los staatva nd emycologie ,doorda td e vraagstelling vegetatiekundig is,maa roo ktamelij klo sstaa tva nd evege - tatiekunde,doorda t eigen -aa npaddestoele n aangepaste- methode nworde n gebruikt.We lzij nbeid especialisaties ,vegetatiekund ee nmycologie ,nodi g ommycosociologisc honderzoe kt ekunne ndoen .W ebeschouwe nd epaddestoelen vegetatie,d emycocoenose ,ne tal sd evegetati eva nvaatplante ne nmossen , de fytocoenose, als eenonderdee lva nd elevensgemeenschap ,d ebiocoenose . De fytocoenoseword t gebruikto m debiocoenos e teherkenne n eni sdu sd e basiswaaro pd epaddestoelenvegetati ebestudeer dwordt .W ebeschrijve ndaar om geen apartepaddestoelenassociaties ,maa rbeschouwe n depaddestoelen vegetatie alsonderdee l vand ebiocoenos e eni nd epraktij kal sonderdee l vand efytocoenose . DoorYpelaa rwa si n197 2e n197 3ee nonderzoe kgedaa nnaa rd epaddestoe len in eikenkreupelbosjes opvoormali g stuifzand,he tDicrano-Quercetum . Indi tvegetatietyp e bleken zeervee l soortenpaddestoele nvoo rt ekomen , waarondervee lzeldzam esoorte ndi e (totda ntoe )nie tva nander evegetatie - typen in Drenthebeken d waren.He tonderzoe k waarvan de resultatenhie r voor u liggen,i sopgeze t om depaddestoelenvegetati e van enkele andere typeneikenbo so pvoedselarme ,zur e zandgrond tebeschrijve ne nt everge lijkenme t dieva nhe tDicrano-Quercetum .Hierto e zijnva nd ei nDrenth e aanwezige eikenbossen 3type ngekozen :1 .he tQuerco-Betuletum ,eiken-ber - kenbos;2 .he tViolo-Quercetun ttypicum ,ee neikenbo swaa rvoora ld ekruid - laagsoortenrijke r is;3 .he tViolo-Quercetu milicetosum ,ee ntyp eeikenbo s met een uitgebreide boom- en/of struiklaag vanhulst .Teven s werdenee n paarproefvlakke nva n hetDicrano-Quercetu m (dieoo ka ldoo rYpelaa rware n onderzocht)i nhe t onderzoekbetrokken ,o mhe tinzich ti nd epaddestoelen vegetatieva nhe tDicrano-Quercetu mverde rt everdiepen . Devegetati eva ngroen eplante ne nmosse nva nd ebestudeerd ebostype ni s ookbeschreven .Di tgebeurd eo md eproefvlakke nt ekarakterisere ne no mn a tekunne n gaanwelk e associaties hetbetrof .Me tbehul pva nd eliteratuu r moesten ook enkele syntaxonomischeprobleme nworde nopgelost .He twa snie t duidelijkwelk enaa mva nhe tQuerco-Betuletu m enva nhe tViolo-Quercetu md e juiste naam was.Bi jhe tViolo-Quercetu m wasbovendie n deplaat s inhe t syntaxonomischesystee monzeker .He tViolo-Quercetu m ilicetosumwa see nno g nieteerde rbeschreve nbostype ,da tdu sbeschreve nmoes tworde ne ningedeel d inhe t systeem vanvegetaties .He tDicrano-Quercetu m was nog nieteerde r uitgebreid beschrevenvoo rNederlan d ene rwa s geenovereenstemmin g over zijnsyntaxonomisch eplaats . Werkwijze Deproefvlakke n werden allegekoze ni nhe t(plantengeografische )Drent s district. Erwerde nva nhe t'Dicrano-Quercetu m 3proefvlakke n uitgezet,8 proefvlakkenva n hetQuerco-Betuletum , 10va nhe tViolo-Quercetu mtypicu m en8 va nhe tViolo-Quercetu m ilicetosum..D eproefvlakke nwerde nme tpaaltje s gemarkeerd, zodat steedsprecie s dezelfdeple k teruggevonden konworden . Dit isnodi gomda tnie tme téé nopnam eva nd epaddestoelenvegetati e inee n gunstigetij do fme tenkel eopname ni néé njaa rka nworde nvolstaan .D ever schijningva npaddestoele n hangtnoga la fva nd eweersgesteldheid ,e rzij n duidelijke seizoensaspecten en ook kand epaddestoelenflor a van jaarto t jaar ergverschillen . Indi tonderzoe kzij nd eopname ngemaak ti nd ezome r enherfs tva n 1976,1977 ,197 8 en1979 .D eopname nzij npe rproefvla kge combineerdto téé nsynthetisch eopname . Bijhe tkieze nva nd eproefvlak-groott ewer duitgegaa nva nd emycosocio - logischevraagstelling .Ho egroo tee nproefvla kmoe tzij no mrepresentatie f tezij nbi jpaddestoelenopnamen ,i snie tbekend .Verschillend eauteur skoze n dieafmetinge ndi ei nhu nvraagstellin ghe tmees tgeschik twas ,e nhanteer den dus elk andere afmetingen,va n 1m 2 totenkel e tientallenhectares . Bepalingenva nhe t 'minimum areaal'zij nmaa rzelde n gedaan.Ypelaa r(i n voorber.)schatt e hetminimu m areaal inhe tDicrano-Quercetu m (onderzocht gedurende 2jaar ) op 600m 2. Jahn,Nespia k enTüxe n (1967)schatte nda t 1000m 2 inhe tLuzulo-Fagetu mleucobryetosu m (4jaa ronderzocht )represen - tatiefza lzijn .Winterhof f (1975)von di nee n Festuca iemani-associatie (eenmaa londerzocht )da t100 0m 2 nietrepresentatief ,du st eklei nwas . Eenproefvla kkieze nw ebi jvoorkeu ri nee nhomogen evegetatie ,hie rdu si n eenhomogen epaddestoelenvegetatie .Aa nee npaddestoelenvegetati e isechte r nietz ogemakkelij kt ezie no fdez ehomogee nis .Daaro mzij nd eproefvlakke n daargekozen ,waa rvegetati eva nhoger eplante ne nmosse ne rhomogee nuit zag.E rzij nechte r (bijna)gee nhomogen evegetatie st evinde nwaari nproef vlakken groterda n100 0m 2 uitt ezette n zijn. Indi tonderzoe k zijnd e proefvlakken bijvoorkeu r 1000m 2 groot gemaakt,ee nafmetin gdi ewaar schijnlijkwe lrepresentatie fi se nno gne tgeschik ti so mbinne nee nrede lijketij d (4-6uur )ee npaddestoelenopnam et emaken .Ee nenkel emaa lmoes t echter eenkleine r oppervlakteworde ngekozen ,i nverban dme td ehomogeni teitva nd efytocoenose .Waa rda tmogelij kwa sbinne nee nhomogen efytocoe - nosewer dnaas the tproefvla kno g500-200 0m 2 geïnventariseerdo ppaddestoe len.Hierbi jwer d alleengele to psoorte ndi ei nhe tproefvla knie tware n gevondene nwer dallee nd eaanwezighei dgenoteerd . Een paddestoelenvegetatie wordt beschreven met 'abundantie opnames', opnames waarbij van elkepaddestoelensoor the taanta l exemplareno pee n bepaaldeoppervlakt eword tgetel do fgeschat ,i nander ewoorde nwaarbi jva n elke soortd eabundanti eva nd evruchtlichame nword tbepaald .Hoewe lnie t alleproefvlakke neve ngroo tzijn ,zij nhie rall eabundanti ewaarden ,even tueeln aomrekening ,betrokke no p100 0m 2. Hetaanta lvruchtlichame ni sd eresultant eva nhe taanta lmyceli ae nva n het aantalvruchtlichame npe rmycelium .D ewerkelijk e abundantieva nee n paddestoelensoort iseigenlij k deabundanti eva nd emycelia .He ti shelaa s onmogelijkdez edirec twaa rt enemen .He tenig ewaarui td eaanwezighei dva n eenmyceliu m blijktzij nd evruchtlichamen .Darimon t (1975)bepaald ehe t aantal 'stations',he taanta lplekke nwaa rd evruchtlichame nva nee nsoor t staan.Ee nstatio nka ndu séé nmyceliu mzijn ,maa roo kee nfragmen tervan , of twee dichtbijee n gelegenmycelia . Indi tonderzoe k werdd egroothei d 'RuimtelijkeFrequentie 'ingevoer do md eabundanti eva nd emyceli at eschat ten.D eRuimtelijk eFrequenti eword tbepaal di nproefvlakke ndi ezij nonder verdeeld indeelproefvlakje sva n2 5m 2 elk.Getel dword ti nhoevee lva nd e deelproefvlakjesee nsoor tvoorkom t ('frequentie opname1).D eRuimtelijk e Frequentieva nee nsoor ti she tdee l (percentage)va nhe taanta ldeelproef vlakjes waar devruchtlichame nva ndi esoor tzij ngevonden .Doo rdez eme thodeko noo kd ehomogenitei tva nd eproefvlakken ,d eruimtelijk everdelin g vansoorte nbinne nee nproefvla ke nd ecorrelati etusse nsoorte nbestudeer d worden. Van alleproefvlakke n zijnopname ngemaak tva nd evegetati eva nvaat - planten eno p de grond groeiendemosse ne nkorstmossen .Apart e opnamen werden gemaakt vanmosse n enkorstmosse ndi eo pdood ,rotten d houtva n stronken, stobbeno fstamme n groeiden,omda tdi tsynusi ahe tsubstraa ti s voorsommig epaddestoelen .I nproefvlakke ndi eware ningerich to mfrequenti e opnamest emaken ,wer dvoo rd eopnam eva nd efytocoenos ehetzelfd eoppervlak te gekozen alsvoo r de opnames vand epaddestoelenvegetatie . Dezeopnam e werdoo kal sfrequenti eopnam egemaakt ,o md efytocoenos eme td emycocoenos e tekunne nvergelijken . Ind eproefvlakke ndi enie tware ningerich to mfre quentie opnames temaken ,wer d voor de opnameva n defytocoenos eee nwa t kleineroppervlakt egekozen ,namelij k150-20 0m 2.D evegetati eopname szij n opd e gebruikelijke wijzei nee ntabe lgerangschikt ;d evolgord ewaari nd e proefvlakkenkwame nt estaa ni soo kbi jd etabelle nva nbode me npaddestoe lenaangehouden . Voor eennader e karakterisering van deproefvlakke nzij noo kd ebodem profielenbestudeerd , enwer d een aantalbodemchemisch ekenmerke nva nd e A0laa g (halfverteer d strooisel)e nva nd eA2 1laa g (zandme tingespeld e humusbepaald . SAMENVATTING In ditproefschrif tword td epaddestoelen-vegetatie , devegetati eva n hogereplanten ,mosse n enkorstmossen ,e nd ebode m (profielene nchemisch e analyses)beschreve nva nd eassociatie sDicrano-Quercetum ,Querco-Betuletu m enViolo-Quercetu me nva ndien ssubassociati eilicetosu mi nDrenthe . Devegetati e vanhoger eplanten ,mosse n enkorstmosse nwer dbeschreve n metopname svolgen sd eBraun-Blanque tmethode ,gewijzig dvolgen sBarkma ne n anderen.He t synusiava nmosse n op rottende stronken, stobbene nstamme n werd apartbeschreven ,omda the the tsubstraa ti svoo rsommig epaddestoel soorten.D e juistenaa mva nd evegetatietype ni sbepaal dme tbehul pva nd e literatuur,evenal sd esynsystematisch eplaats .D ei nd eliteratuu rgenoem deken -e ndifferentiërend e soortenzij nbesproken .He tQuerco-Betuletu me n hetViolo-Quercetu m blekennau wverwant e associaties tezijn .Z ebehore n tot het verbond Quercion robori-petraeae.He tViolo-Quercetu m ilicetosum werd alsnieuw e subassosiatiebeschreven .D emenin gva nBarkman ,da the t Dicranoi-Quercetum een zelfstandige associatie is,ko nworde nbevestigd . Deze associatie heeftvee l differentiërende soorten (vrijweluitsluiten d mossen en korstmossen), zodat de verwantschap methe tQuercio nrobori - petraeaegerin gis .He tword tdaaro mnie tto tdi tverbon dgerekend . Deonderzocht eproefvlakke nva nhe tQuercio nrobori-petraea ehadde nall e eenzandig e boden,vaa kme tee nondergron dva nlee mo flemi gzand .I nee n proefvlak was eenbroe k eerdgrond aanwezig,d eander eproefvlakke nhadde n alleduidelijk e humuspodzo lprofielen .D eproefvlakke ni nhe tViolo-Quer cetum ilicetosum hadden relatiefdiep eprofiele nwaari nvaa kee ndikk eAl h laag aanwezig was.I nhe tViolo-Quercetu m typicume nhe tQuerco-Betuletu m was hetprofie l korter enwa s eenAl hlaa g gewoonlijk niet aanwezig.D e bovenste laagva n debode mva nd eQuerco-Betuletu mproefvlakke nblee kove r het algemeen gestoord tezijn .D eproefvlakke n vanhe tDicrano-Quercetu m bleken te liggen optamelij k recent stuifzand waarin slechts eenuiters t kortprofie lwa sontwikkel d (duinvaaggron dme tee nmicropodzol) . De chemische analyses werden gedaan inmonster sva n deA O en deA2 1 laag. Indez e lagen ishe t eestemyceliu m aanwezig.D ebodem sbleke nall e zeerzuu rt ezijn ,p H2, 9 -3, 5 ind eA2 1laag ,3, 3 -4, 7 ind eA Olaag .I n deA O laagware n deNa ,K ,Mg ,e n Caionene n (uitgedrukti nmassafracti e enmassaconcentratie )i nel kva nd evegetatietype ni nnagenoe ggelijk ehoe veelheden aanwezigt e zijn,i nd eA2 1laa gware nd ehoogst econcentratie s inhe tViolo-Quercetu m ilicetosum en de laagstei nhe tDicrano-Quercetum . De 'totaleione nvoorraad 'wa she thoogs ti nhe tViolo-Quercetu m ilicetosum, veel lager inhe tViolo-Quercetu m typicum enhe tQuerco-Betuletum ,e nno g weer lager inhe tDicrano-Quercetum . DeC/ Nverhoudin ggedroe gzic homge keerde nna mto ei ndez evolgorde . Depaddestoelen-vegetati e isbeschreve nme tbehul pva nsynthetisch eop namenva nproefvlakke ndi egedurend e4 jaa ronderzoch tzijn .E rwerde nvee l meersoorte npaddestoele n (314)da ngroen eplante n (72)o fmosse ne nliche - nen (67)gevonden .Criteri ao mdifferentiërend e soortenbi jfung it eonder scheidenwerde ngeformuleerd . HetDicrano-Quercetu m enhe tQuercio nrobori-petraea ehebbe nte nopzicht e vanelkaa rvee l differentiërende soorten.Di tondersteun the tide eda the t Dicrano-Quercetumnie tto the tQuercio nrobori-petraea ebehoort .He tQuerco - Betuletume nhe tViolo-Quercetu mhebbe nte nopzicht eva nelkaa rminde rdif - ferntiërendesoorten ,maa rzij nwe lduidelij kgescheiden .Zowe lhe tDicrano - Quercetum als het Violo-Quercetum ilicetosum,beid e vegetaties metzee r weinigvaatplanten ,bleke nme tbehul pva n depaddestoelen-vegetati ebete r tekarakteriseren .Di t geldtme tnam evoo rhe tViolo-Quercetu m ilicetosum, datoo kzee rar mi saa n (terrestrische)mosse ne nkorstmossen . Het 'minimum areaal',o feigenlij kd erepresentatiev eproefvla kgroott e bleeki nd eassociatie sva nhe tQuercio n1.50 0m 2 tezijn ,e ni nhe tDicra no-Quercetumminsten s3.00 0m 2. Omd ewerkelijk eabundanti eva nfungi ,du sd eabundanti eva nd emycelia , tebepale n werd eennieuw egroothei dingevoerd ,d eTotal eRuimtelijk eFre quentie (TotalSpatia lFrequency ,TSF) :he tdee l (percentage)va nhe taanta l deeïproefvlakkenva n eenproefvla k waarin ineni gjaa rvruchtlichame nva n die soortzij nwaargenomen .D e absolutemaximal eabundanti e (AMAC)va nd e vruchtlichamen vertoontwe l eenbepaal d verbandme td eTSF ,maa rhe ti s gewoonlijknie tmogelij kd een eeenhei di nd eander eo mt erekenen . Debepalin gva nd eTS Fga foo kee naanta lander eresultaten .Z oko nwor denvastgestel d dat dederd ewe tva nRaunkae roo kopgaa tvoo rd efung iva n hetQuercio nrobori-petraeae ,maa rnie tvoo rd efung iva nhe tDicrano-Quer cetum,di ti ntegenstellin gto td egroen eplante nva nhe tDicrano-Quercetum . De fytocoenose vanhe tDicrano-Quercetu mblee kdoo rhe tvoorkome nva nuit gestrekte mos-tapijten eenkarakteristiek e heterogeniteit tebezitte ndi e groteri sda nbi jd eander esyntaxa . Ook kondenverspreidingspatrone n vansoorte nbinne nd eproefvlakke nbestu deerdworden :verui td emeest esoorte nbleke nnormaa lversprei dt ezij ne n erware ngee nsoorte ndi eregelmati gversprei d (onderdispers)waren .Slecht s 35soorte nware n in2 o fmee rproefvlakke n geklusterd (overdispers).Di t kanworde nveroorzaak t doordatd emyceli a zogroo tzij nda tzi jove rver scheidenedeeïproefvlakke nversprei dzijn ,o fdoorda td emyceli agekluster d groeien,bijvoorbeel ddoorda the tproefvla kinhomogee nis ,althan svoo rdi é soort. Ookkonde ndankzi jdez emethod ecorrelatie stusse nverspreidingen ,zowe l van fungi onderling alsva n fungime tvaatplante no fmosse nworde nbestu deerd. Inbeid e Dicrano-Quercetum proefvlakken bleek een correlatiet e bestaan tussenverscheiden e paddestoelsoorten (bv. Marasmius androsaceus, Galerina decipiens, Inocybe napipes) end emos-tapijten .I nee nViolo-Quer cetumtypicu mproefvla kbeston dee ncorrelati etusse nee nhoo gaanta lpadde stoelsoortene nee nlag ebedekkin gva nd ekruidlaag . Marasmius epiphylloides correleerde metterrestrisc h groeiendeklimop ,maa rkwa mnie tvoo ri nall e deelproefvlakkenme tee nhog ebedekkin gva nd eklimop . Collybia cookei cor releerde in een aantal proefvlakkenme t Armillariella mellea. Daarnaast zijnno genkel eander ecorrelatie sbesproken . Depaddestoele n werden ingedeeld naar oecologische groepen (waarvand e mycorrhizasoorten ,d ehumus-saprofyten ,d esaprofyte no phou te ntakke ne n de saprofyten opblad ,vruchten ,plante ne nmosse nd ebelangrijkst ebleke n tezijn )e nhe taandee lva nel kva nd egroepe ni nd epaddestoelenflor awer d bepaald. Inhe tDicrano-Quercetu mbleke nzee rveel ,i nhe tViolo-Quercetu m ilicetosumweini gmycorrhiz a soortenvoo r tekomen .He tDicrano-Quercetu m hadweini g saprofyten ophout ,he tViolo-Quercetu m ilicetosumweini ghumu s saprofyten.D e saprofyten opbla d e.d. warenhe ttalrijks ti nhe tQuerco - Betuletume nhe tViolo-Quercetu m typicum. Van een aantal paddestoelensoorten, namelijk diewaa r de determinatie moeilijkhedengaf ,doo rd eaar dva nhe tmateriaa lo fdoorda te rtaxonomisch e problemenwaren ,zij nkort eto tlang eaantekeninge no fbeschrijvinge nopge nomen. Psathyrella fulvescens var. dicrani werd alsnieuw evariëtei tbe schreven. 1 INTRODUCTION TheBiologica lStatio n (provinceo fDrenthe ,Th eNetherlands) ,a sectio n ofth eAgricultura l University Wageningen (TheNetherlands) , hasfo rman y yearsbee na centr eo fresearc ho nth esociolog yan decolog yo fmacrofungi . Research hadbee ndon ei nJunipe rscrub s (Barkmanan dD eVries) ,i ngrass landvegetation s (Arnolds), inPice awoods ,Callun aheathers ,oa kscrub s (otherco-worker san dstudents) . Indescribin gth efungu svegetatio no fal l phytocoenoses inth eDrenthia ndistrict ,i ti shope dt oaquir ea ful lknow ledge on the sociological range and status,an d ecological range ofal l macromycetesi nDrenthe .Thi sknowledg ei sals oexpecte dt ohel psolv eman y synsystematicalproblems ,especiall y inassociation swit hfe wphanerograms , mosseso rlichens . Onlymacromycete s areinvolve d inthes e studies:al lHymenomycete san d Gasteromycetes, and those Ascomycetes and Aphyllophorales with a well developed, not very small fruiting body.D eVrie s is also studying the smallAscomycete s andAphyllophorale s ofth eJunipe rscrubs . The study ofth efungu s'vegetation srequire sit sow nmethods ,suite dt o thefungi .I ti sno tpossibl et ous eexactl yth esam emethod sa sstudyin ga phytocoenose.W econside rth efungu svegetation ,th emycocoenose ,a sa par t ofth ebiocoenose , justlik eth ephytocoenos e isa par to fth ebiocoenose . Thephytocoenos ewa suse dt orecognis eth ebiocoenose ,an ds oform sa basi s for the study of themycocoenose .W e therefore do notdescrib eseparate , independentfungu svegetations . The fungusvegetatio n ofoa k scrubs on former driftsand, theDicrano - Quercetum (Ypelaari nprep. )appeare dt ob ever yspecie sric han dt ocontai n manyrar especie sknow nonl yfro mthi styp eo fvegetation . The object of this thesiswa s to describe the fungusvegetation s of other types ofoakwoo d onnutrien tpoor ,acid ,sand ysoil ,an dt ocompar e itwit h thato fth eDicrano-Quercetum .Thre etype so foakwoo dwer echosen : 1.Querco-Betuletum ,oak-birc hwood ;2 .th eViolo-Quercetu mtypicum ,a typ e witha mor evarie dherblayer ,an d3 .th eViolo-Quercetu m ilicetosum,a typ e with awel l developed holly (Ilex aquifolium)tre e or shrublayer .Thre e Dicrano-Quercetum plots (already investigatedb yYpelaa ri n197 2an d1973 ) were included, in order toge tmor e detailed information of the fungus vegetationo fth eDicrano-Quercetum . Thevegetatio n ofphanerogams ,mosse san dlichen swa sals odescribe dt o characterizeth eplots ,t odetermin eth enam eo fth esyntaxon ,an dt osolv e some synsystematical problems.Th e correctname s forth eQuerco-Betuletu m andth eViolo-Quercetu m hadt ob edetermined .Th esynsystematica lpositio n of theViolo-Quercetu mwa suncertain .Th eViolo-Quercetu m ilicetosum had notpreviousl ybee ndescribed ,thi swa sdone ,togethe rwit ha synsystemati - calclassification . Inth eNeterland s theDicrano-Quercetu m had notbee n described extensively and therewa s no agreemento n its synsystematical position. All plots were situated in theplantgeographica l districto f Drenthe.Thre eplot s weremad e in theDicrano-Quercetum ,8 i nth eQuerco - Betuletum,1 0i nth eViolo-Quercetu mtypicum ,an d8 i nth eViolo-Quercetu m ilicetosum. Theplot swer efixe dwit hpicket si norde rmak ei tpossibl et o refind exactly the sameplace .Thi swa snecessar ybecaus eon erelev éi na good season or severalrelevé si non eyea ronl yi sno tsufficien ti norde r tob eabl et odescrib ea fungu svegetation ,a sth eformatio no fcarpophore s dependsgreatl yo nth eweather .Distinc tseasona laspect sexis tan dth efun gusflor aca nals odiffe rmuc hfro myea rt oyear .I nthi sstud yth erelevé s weremad ei nth esumme ran dautum no f1976 ,1977 ,197 8an d1979 .Th erelevé s werecombine dpe rplo tint oon esynthetica lrelevé . The size ofth eplot s depended onth e sizeneede dfo rmakin ga fungu s relevé,bu tth eplo t areatha t isrepresentativ e fora relev éo fa fungu s vegetation, isunknown .Author s haveuse d different areas,sui tt othei r own situation, sousin g areas of1 m 2 upt ote no rmor ehectares .Deter mination ofth e 'minimalarea 'ha sbee n done rarely.Ypelaa r (inprep. ) estimated therepresentativ e plotare ai nth eDicrano-Quercetu m (a2 year s study)t ob e 600m 2. Jahn,Nespia k andTüxe n (1967)estimate d aplo to f 1000m 2 in the Luzulo-Fagetum leucobryetosum (research that lasted at least4 years )t ob e representative.Winterhof f (1975)foun di na Festuca lemani-vegetation (one survey)tha t aplo to f100 0m 2wa sno trepresenta tive. A plot ispreferabl ychose ni na homogeneou svegetation ,an dfo ra fun gusrelev é in ahomogeneou s fungusvegetation .A shomogeneit yo fa fungu s vegetation isno teasil ysee nth eplot swer echose ni na phytocoenos etha t looked homogeneous.Homogeneou s phytocoenosesi nwhic hplot so f100 0m 2o r larger canb e situated,wer e sparse. Inthi sstud ya plo tare ao f100 0m 2 was chosen forth erelevé so fth efungu svegetation .Thi sare ai sprobabl y representative andha s the sizet ob eabl et omak ea fungu srelev éi n4- 6 hours.A fewtime s asmalle rplo tha dt ob echosen ,dependin go nth esiz e ofth ephytocoenose .A n areao f500-200 0m 2 around theplo ti nth esam e phytocoenose (depending on thepossibilitie swithi nth ehomogeneou sphyto coenose)wa s alsoinvestigate d forspecie s thatwer eno tpresen t inth e p\otarea .Onl yth epresenc eo fthes especie swa snoted . Inth erelevé so fth efungu svegetatio nth eabundanc eo fth ecarpophore s wa^sdetermine d by counting orestimatin g thenumbe r ofcarpophore si na n areao f100 0m 2. Inplot slarge ro rsmalle rtha n100 0m 2 abundancewa scon vertedan dals ocounte di nnumbe ro fcarpophore spe r100 0m 2. Thenumbe r ofcarpophore s isth eproduc to fth e numbero fmyceli aan d ienumbe r ofcarpophore spe rmycelium .Th erea labundanc ei si nfac tth e 10 abundance ofmycelia ,whic hcanno tb edirectl ydetermined .Darimon t (1975) determined thenumbe ro f 'stations',th enumbe ro fpatche swher eth ecar pophores ofa specie s occurred.A stationca nb eon emycelium ,a fragmen t ofa mycelium ,o r2 myceli agrowin gclos et oeac hother .I nthi sstud yth e value 'SpatialFrequency 'wa screate dt oestimat eth eabundanc eo fmycelia . The SpatialFrequenc y isdetermine di nplot sdivide dint opartia lplot so f 25m 2.Th enumbe ro fpartia l plotswher e aspecie s occurredwer ecounte d ('frequencyrelevé') .Th eSpatia lFrequenc yo fa specie si sth eproportio n ofth epartia lplot swher eth ecarpophore so ftha tspecie swer efound .Thi s methodwa sals ouse dfo rth edeterminatio no restimatio no fth ehomogeneit y ofth eplots ,o fth espatia ldistributio no fth especie si nth eplots ,an d ofth ecorrelatio nbetwee nspecies . Relevés were made ofth evegetatio n ofphanerogams ,an dterrestria l mosses andlichen s inal lplots .Separat erelevé swer emad eo fth emosse s and lichens growingo ndead ,rotte nwoo do ftruncs ,stumps ,an dlogs ,be causethi ssynusi ai sth esubstrat efo rsom efungu sspecies .Th erelevé so f thevegetatio no fhighe rplant s inplot suse dfo rfrequenc yrelevés ,wer e takena slarg ea sth erelevé so fth efungu svegetations ,i norde rt ob eabl e tocompar e thefungu svegetatio nwit hth evegetatio no fhighe rplants .I n plots thatwer eno tuse d forfrequenc yrelevés ,a smalle rare awa schose n for the relevés ofth ephytocoenose : 150-200m 2.Th evegetatio n relevés werearrange di na tabl ei nth eusua lway .Th esequence so fplot swer eals o usedi nth etable so fth esoi lcharacteristic san dth efungi . The soil profiles ofth eplot swer e studied, andsom esoi lchemica l factors of theA O (fermentation layer)an dth eA2 1laye r (mineral layer with infiltrated humus)wer edetermine di norde rt ocharacteriz eth eplot s further. 11 2 THE PHYTOCOENOSES 2.1 INTRODUCTIONAN DMETHOD S Theplot swer e selectedi nhomogeneous-lookin g vegetations.Th esiz ewa s at least15 0 m2, whichconforme dt oth erepresentativ e plotare afo r vege tationi nwoods .Al lplots ,however , were larger,ca .100 0m 2 (exceptfo r some plotso fth eViolo - Quercetum ilicetosum, ifa large r areawa s not available.Se eplo t areas intabl e 1). Anare ao f100 0m 2wa sno tneede d to describeth evegetation ,bu tt odescrib eth efungu svegetation .Fo r1 7 plots thewhol e plot (1000m 2)wa sdescribe dan dno tonl ya representativ e parto fth eplot .Relevé so f15 0o r10 0 m2 andth emos slaye ro fal lrelevé s (excepti nth eplot s2 an d3) , weremad ea son erelevé ,thos eo f875 , 1000, or 1050m 2wer e madei npartia l relevéso f2 5m 2, comparablewit hth efre - guencyrelevé sfo rth efungu svegetation .Partia l relevéswer e laterunite d toon e'total 'relevé ,usin gth evalue s givenbelo wfo rMea nRea lCover . The vegetations were described with Braun-Blanquet relevés.Th ecove r values aregive ni nBraun-Blanque t symbols -modified accordingt oBarkma n et al. (1964)- andar eexplane d asfollow s (mentioned inparenthese sar e thevalue sfo rth e'Rea lMea n Cover'): r rare,1- 2specimen s (0.1%) + ' littlenumerous ,3-1 0specimen s (0.5%)^CQVe r always ^5 % 1 numerous,10-10 0specimen s(1% ) 2m verynumerous ,> 10 0specimen s(3% ) 2a cover 5 -12.5 %( 8% ) 2b cover 12.5-2 5 % (18%) 3 cover2 5 -5 0 %(35% ) 4 cover5 0 -7 5 % (60%) 5 cover7 5 -100% (86%) In some plots arathe r large areawa scovere db ydead ,hal frotte nwoo d of stumps, stubs, trunks,an dfalle n logs, often witha remarkabl y high mosscover. Invegetatio n relevés this non-terrestrial synusiai susuall y not included,bu ti nthi s study this synusiawa simportant ,becaus e some fungus species only occured onthi s substrate. 'Stump relevés' therefore were made inadditio nt oth evegetatio n relevé. Firstly anestimat ewa s madeo fho wmuc ho fth eplo t areawa scovere db ydea d stumpsan dlogs ,the n a separate relevéwa smad eo fthi ssynusia .Th ecove ro fth emos s layeran d of themos s specieswa sestimate d asa portio no fth eare ao fth edea d stumps andlogs . These relevés were made justa sthos eo fth eterrestria l moss layer,namel ya son erelev épe rplo t (noti npartia lrelevé sa si nth e 12 largeplots) . Thevegetatio n relevéswer e made in the summer of 1977an d 1978.Th e relevés ofth e terrestrialmos slaye ran do fth emosse so ndea dstump san d logswer emad ei nMa y197 9togethe rwit hJ.J .Barkman .Th erelev éo fplo t1 hasbee ncopie d fromVreugdenhi l& Barkma n (ReportBiol .Station ,Wijster) . Inth esynoptica ltabl e (table2 )th ePresence ,P ,i na scal eo f1 0part s inRoma nnumerals ,an d theTota l CoverValue ,TCV , aregiven .Th eTC Vi s themea ncove rx 100 ,calculate d with thevalue s forth eRea lMea nCove r (see above). The synopticaltabl ewa scalculate d forth eDicrano-Quercetu m from 42relevé s givenb yVreugdenhi l& Barkman ,thos efo rth eothe r (sub)- associations fromth erelevé sgive ni ntabl e1 . Therear emethod sfo rcalculatin gmathematica laffinit ybetwee n2 associ ations.I nthi spape rBarkman' smethod s (1974)wer eused .A isth eaffinit y based onth enumbe r ofcommo n and not common species,an dwa scalculate d accordingt o i5 Ac=(c+l)/[(ab) +1 ] (ci sth enumbe ro fspecie si ncommo ni nbot hassociations ,a i sth enumbe r of species occurring only in theone ,b the numberoccurrin gonl yi nth e other asociation).Therefor e onlyabsenc ean dpresenc eo fspecie swa suse d tocalculat eA . Basedo nth evalue sfo rPresence ,P ,i sth eaffinit y 15 Ap=ZCp/(XAp2Bp) (C_i sth ePresenc evalu ei ncommon ,s oth elowes to fth etw oP value so fa species,A _ isth e Presencevalu e ofa specie slessene dwit hC p infavou r ofth eon eassociation ,B _th esam efo rth eothe rassociation) . Basedo nth ecove rvalue so fth especies ,TCV ,i sth eaffinit y AT=ICT/(IAT2BT)^ (CTi sth eTC Vvalu ei ncommon ,s oth elowes to fth e2 TC Vvalue so fa spe cies,A _i sth eTC Vo fa specie slessene dwit hC _i nfavou ro fth eon easso ciation,B ™ the same in favour of the other association). So forth eA _ bothth evalue s forpresenc ea swel la sthos efo rcove rwer eused . 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XI S S •• -H TJ 0) 3 • ? 01 OIT) •d u ••ois.g XI 0)—J< QXI« -H <0 33-H E •H • tn o 0) oi "O U 03 • M 0) O) O 01 — U S H ? — -asa. c UJ>—OiO-^S S3 ro ca Q m •0 w -»—.--i M- 10 rH • >-^M^ U M s Annext otable s1 an d2 . Thefollowing ,rar especie sar eno tmentione di nbot htables .Mentione dar e thecove rvalu e inBraun -Blanque t symbols and, inparentheses ,Presenc e andTota l CoverValue . Ifno t indicatedotherwis ei nth emos sspecies ,i t wasmos stha twa sgrowin gterrestrially . Plot1 : Rumex acetosella L. r(I-l), Nardus stricta L.r(I-l), Diplophyllum albicans (L.) Dum. +(1-1), Buxbaumia aphylla Hedw. +(1-1), Calypogeia trichomanis (L.)Cord a +(1-1), Ceratodon purpereus (Hedw.)Brid . +(1-1), Cetraria glauca (L.)Ach . r(II-4). Plot2 : Juncus squarrosus L. r(I-l), Polytrichum piliferum Hedw. r(I-4), Cladonia gracilis (L.)Willd . +(1-1), Cladonia destricta (Nyl.)Sandst . +(1-1), Cladonia floerkeana (Fr.)Sommerf . r(II-2), Cornicularia aculeata (Schreb.)Ach .+(II-5) , Pinus sylvestris L.treelaye r 1(1-2). Plot3 : Campylopus introflexus Brid.+(1-4) . Plot4 : Polypodium vulgare L.r(I-l), Pleurozium schreberi (Brid.)Mitt ,o n deadstump s2b(1-225) . Plot7 : Lolium perenne L.r(I-l), Chenopodium album L.r(I-l) . Plot8 : Anthoxanthum odoratum L.2m(I-37) , Poa trivialis L.1(1-12) , Luzula cf campestris (L.) DC . +(1-6), Potentilla erecta (L.)Räusche l r(I-l), Prunus spec, herblayer +(1-6), Cladonia squamosa (Scop.)Hoffm . ondea d stumps r(I-l). Plot9 : Isopterygium seligeri (Brid)Dix . 1(1-12), Isopterygium elegans (Hook)Lindb .o ndea dstump s r(I-l). Plot11 : Plagiothecium undulatum (Hedw.)B.S.G .+(1-6) . Plot12 : Moehringia trinervia (L.) Clairv . 1(1-10), Senecio sylvaticus L. +(1-5), Poa pratensis L.r(I-l), Veronica officinalis L.+(1-5) . Plot13 : Dicranum majus Sm.o ndea dstump s r(I-l). Plot14 : Dicranum fuscescens Turn,o ndea dstump sr(I-l) . Plot15 : Plagiothecium latebricola B.S.G.+(1-5) . Plot17 : Holcus mollis L. +(1-5). Plot22 : Hedera helix L.i ntre elaye rr(I-l) . Plot25 : Dactylis glomerata L. r(I-l), Poa annua L. r(I-l), Polygonum aviculare L.+(1-6) . Plot26 : Polytrichum marginatum Web.e tMohr ,o ndea dstump sr(I-l) . Plot27 : Sambucus nigra L.her blaye r+(1-6), Bryum capillare Hedw.o ndea d stumpsr(I-l) . Plot28 : Polygonum cf convolvulus L.r(I-l) . Plot29 : Acer pseudoplatanus L. shrublaye r r(I-l), Anemone nemorosa L. r(I-l), Corylus avellana L. herblaye rr(I-l), Taxus baccata L. (seedling) r(I-l). Notmentione di ntabl e2 ar esom especie swit hPresenc eI (mentione di npa renthesesi sth eTota lCove rValue) , inth eDicrano-Quercetum : Orthodicranum montanum (Hedw.)Loesk . (5),Dicranum fuscescens Turn.(1) ,Dicranum majus Sm. (7);i nth eViolo-Quercetu m ilicetoscum: Fagus sylvatica L. inshru b layer(1) . 21 2.2 RESULTS Theresult sar egive ni na vegetatio ntabl e (table1 )an di na synoptica l vegetation table (table 2). Thevegetation sar edistinguishe di nth eusua l way into tree,shrub ,her b andmos s layers. Inth etable sth elayer sar e not given separately. The layer fromwhic ha specie scam ei salway snote d before the species,wit ht (tree layer),s (shru blayer) ,h (herblayer) , andm (moss layer). The relevés ofth emosse so ndea dstump san dlog sar e givenseparatel ya tth een do fth etables . Theassociation sar ediscusse dseparately . 2.2.1 The Querco-Betuletum Quercorobori s- Betuletu mTx .1930 . Type relevé Tüxen 1930,p .7-8 ,relev é2 ,a sQuerceto-Betuletum .Fo rgram maticalreason sth enam eha st ob eQuerco-Betuletum . Synonyms: Molinio- Quercetu mrobori sScam ,e tPass .1959 . Vaccinio- Quercetu mloniceretosu mDoin g1962 . Melampyro- Quercetu mrobori sPass ,e tHofm .1968 . Westhoff& De nHel d (1969), who reviewedth eQuerco-Betuletu mincludin g theDicrano-Quercetu m (tob ediscusse dfurthe ron )liste dth efollowin go f differential taxa in relation toth eViolo-Quercetum : Pinus sylvestris, Polypodium vulgare, Vacciniuiû vitis-idaea, Calluna vulgaris, Festuca tenuifolia, Juniperus communis, Empetrum nigrum, Cladonia sect. Cladina, Dicranum scoparium, Aulacomnium androgynum, Buxbaumia aphylla, Pleurozium schreberi, Leucobryum glaucum, Ptilidium ciliare, Campylopus flexuosus, Polytrichum juniperinum and Rhytidiadelphus squarrosus. Cornus suecica is considered tob e theonl ycharacte rspecie si nth eNetherlands .Oberdorfe r etal . (1967)calle d theQuerc o roboris-Betuletum aweakl y characterized associationpoo r incharacte r species ("Rumpf-association").The ydi dno t namean ydifferentia lspecies .Passarg e& Hofman n (J.968)di dno tnam echar acter or differential species,bu tthe y give asynthetica l table.Doin g (1962)an dBakke r (1969)gav ecertai ngroup so fspecie sa scharacteristic s forthi sassociation . Tüxen(1937 )divide dth eQuerc oroboris-Betuletu mint o2 subassociations : molinietosum and typicum. Theplot s investigated here allebelon g toth e subassociationtypicum . From this survey the following species appeared to be differential species for theQuerco-Betuletu m (seetable s1 an d 2), inrelatio nt oth e Dicrano-Quercetuman dth eViolo-Quercetum : Vaccinium myrtillus, V. vitis- idaea, and Melampyrum pratense; forth eQuerco-Betuletu m and theViolo - Quercetum together inrelatio n to theDicrano-Quercetum : Molinia caerulea 22 and Rubus idaeus; for theQuerco-Betuletu m and theDicrano-Quercetu m to getheri nrelatio nt oth eViolo-Quercetum : Leucobryum glaucum and Campylopus flexuosus. Iti sclea rtha tsom etax acalle ddifferentia ltax ab yWesthof f & DenHeld , are inm y opinion differentialtax afo rth eDicrano-Quercetu m inrelatio n to theQuerco-Betuletu m and theViolo-Quercetum , and notfo r theQuerco-Betuletu m inr.elatio nt oth eViolo-Quercetum . Itconcern s Calluna vulgaris, Festuca tenuifolia (=F. ovina var. tenuifolia), Empetrum nigrum, Cladonia sect. Cladina, Dicranum scoparium, Aulacomnium androgynum, Pleuro- zium schreberi, Ptilidium ciliare, and Campylopus flexuosus. Theothe r species they mentioned can not be treated asdifferentia l or character species: Pinus sylvestris, Juniperus communis, Polypodium vulgare, Buxbaumia aphylla, Polytrichum juniperinum, Rhytidiadelphus squarrosus, and Cornus suecica. Ionl y found Pinus sylvestris veryrarel yi nth eDicrano-Quercetum , and never in the Querco-Betuletum. This species will notstar t togro w spontaneously in awood , neitherwil l Juniperus communis, whichwa sno t found in the vegetations investigated. Polypodium vulgare and Buxbaumia aphylla were observed only once inth eQuerco-Betuletum , andth eDicrano - Quercetum respectively socanno tb etreate da sdifferentia lspecies . Poly trichum juniperinum, Rhytidiadelphus squarrosus, and Cornus suecica were notobserve d inan yo f thevegetation s investigated. Iti sunlikel ytha t Rhytidiadelphus squarrosus occursi nth eDicrano-Quercetu m ori nth eQuerco - Betuletum; other Rhytidiadelphus species were alsono tobserved . Cornus suecica, aver y rare species and thereforeonl ya loca lcharacte rspecie s. in theNetherlands , ismor e typical fornort h facingwood-edges ,an di s more likely tob e found inassociatio nwit h openwood s thanwit h awel l developedQuerco-Betuletum . From the relevés ofmosse so ndea dstump san dlogs ,n ospecie sappeare d tob e differential forth eQuerco-Betuletu m inrelatio nt oth eothe rasso ciations. There are 8specie s (group7 ) differential for the Querco- Betuletum and the Violo-Quercetum typicum together in relation to the Dicrano-Quercetum: Mnium hornum, Dicranum scoparium, Tetraphis pellucida, Orthodontium lineare, Cladonia glauca, C. digitata, Polytrichum formosum, Plagiothecium curvifolium, and Campylopus flexuosus. Differential species forth eQuerco-Betuletum , andth eViolo-Quercetu m typicuman dilicetosu mi n relation to theDicrano-Quercetu m are Brachythecium rutabulum, Isothecium myosuroides, Lepidozia reptans, and Isopterygium seligeri. The investigated vegetations were scrubbywoods ,coppices ,hig hwoods , or formercoppice s transformed intohig hwoods .I nscrubb ywood san dcop pices the tree layer isno tver yhigh ,an dth eshru blaye ri spoo ro run developed.Usuall y the shrub layer canno tb edistinguishe d asa separat e layer.I nhig hwood sth etre elaye ri shighe ran dmor edistinctl yseparate d from the shrub layer,whic h isbette rdeveloped .Th eher blaye ri snearl y alwayswel l developed.Th e aspect isdetermine d by Deschampsia flexuasa, Holinia caerulea, Corydalis claviculata, and Vaccinium myrtillus, thelas t mentionedsometime scover su pt o65% . 23 2.2.2 The Violo-Quercetum Violo-QuercetumOberd .1957 . Synonyms: Querceto sessiliflorae-Betuletum violetosum rivinianaeTx .e tDiem ,i nTx . 1937 (thisnam e is illegitimate,becaus e thenam e ofth eassociatio nwa s notvalidl ypublished ,Tx .1937) . Fago-Quercetumpetraea e (Tx.1937 )Tx .195 5p.p . Maianthemo-QuercetumBakke r1969 . This association canb e considered asth e form ofth e Fago-Quercetum petraeae in the low-lying plains, in which Quercus robur, and not Q. petraea, dominates (seeVa nde nBroe k& Diemon t1966) . Ifw econside rthi s vegetation as anassociation ,a sha sbee ndon ehere ,th ecorrec tnam ei s Violo-Quercetum. Tüxen (1937), however, considered thisvegetatio n asa subassociation;then ,th ecorrec tnam ei sFago-Quercetu mpetraea evioletosu m rivinianae (Tx.e tDiem ,i nTx .1937 )nov .comb . Doing (1962)distinguishe di nth eNetherland sa tleas t3 vicariou sasso ciations, the Solidagino-Quercetum (=Fago-Quercetu m petraeae (Tx.1937 ) Tx. 1955p.p. )o nacid ,undisturbe dloa msoils ,i nth eSout ho fth eNether lands; theViolo-Quercetu m roboriso nacid ,bu tno tver ypoo rsand ysoils ; the Convallario-Quercetum roboris on sandy soils inth einne rare ao fth e coastal dunes;an dpossibl y afourt h association ofwood swit h Corydalis claviculata and Ilex aquifolium onsoil swit ha loam ysubsoi li nth eNort h East Netherlands. The woods Iinvestigated ,belon g to thisassociation . Bakker (1969)becaus eo fth ever ysligh tdifference scombine dthes e3 asso ciations,int oth eMaianthemo-Quercetu m anddistinguishe d 2subassociations , anemonetosum and typicum. Thewood sI investigated ,belon gt oth esubasso ciationtypicum . Onlya smal lpar to fth echaracte ran ddifferentia lspecie smentione di n literature (e.g.Tüxe n 1937,Oberdorfe r 1957,Oberdorfe r et al.1967 , Westhoff& De nHel d 1969)wa s found. Quercus petraea, Hieracium sabaudum, Luzula luzuloides, and L. sylvatica, mentionedb yWesthof f& De nHeld ,d o not occur inth eDrenthia n district; Rubus saxatilis occurred onlynea r TerApel ; Solidago virgaurea didno toccu ri nth ewoods . Lathyrus montanus wasfoun dmainl yo nth eHondsru gan dth eHavelterberg . Malus sylvestris was found in the Mantingerbos. Populus tremula was found numerous, and Polygonatum verticillatum wasrar ean dsinc edisappeared ,i nth eThijnsbos . These3 specie sdi dno toccu ri nm ysampl eplots .I di dfin d Populus tremula, but itwa s rare,i n 2plot s ofth eQuerco-Betuletum . Hieracium lachenalii was not found inth e investigated plots. Ilex aquifolium is acharacte r speciesi fonl yth etre ean dshru blaye rwer econsidered ;i nth eher blaye r this species occured as often inth eViolo-Quercetu m typicum as inth e Querco-Betuletum. 24 Westerhoff& De nHel d gave alo to fdifferentia lspecie s forth eViolo - Quercetum inrelatio n toth eQuerco-Betuletum . Someo fthes edi tno toccu r oronl yver y rarelyi nth eViolo-Quercetu man dthe nonl yi naberrant ,dis turbedphytocoenoses : Scrophularia nodosa, Narcissus narcissus, Sambucus racemosa, and Selinum carvifolium. Mespilus germanica and Hypericum pulchrum were mainly found in the Southpar t ofth eprocinc e of Limburg (inth e Solidagino-Quercetum). Stellaria nemorum ssp. glochidisperma occurred in Drenthe only in theNorgerholt ,mainl y inth e Stellario-Carpinetum.Al l these species and Viola riviniana were not found inth eplot s Iinvesti gated.Th e other species theymentione d are: Anthoxantum odoratum, Poly- gonatum multiflorum, Luzula pilosa, Atrichum undulatum, Hedera helix, Oxalis acetosella, Convallaria majalis, Stellaria holostea, Corylus avellana, and Prunus serotina. Inthi s survey Anthoxantum odoratum wasfoun donl yonce , ina Querco-Betuletu m ata slightly disturbedspot ;s oi tca nno tb econ sidereda sa differentia lspecies . Oxalis acetosella, Hedera helix, Con vallaria majalis, and Corylus avellana appearedt ob edistinctl ydifferen tial species forth eViolo-Quercetu m (typicum andilicetosu mtogether )i n relation to the Querco-Betuletum and the Dicrano-Quercetum. Stellaria holostea isa differentia lspecie sfo rth eViolo-Quercetu mtypicu mi nrela tiont oth eQuerco-Betuletum ,th eDicrano-Quercetu m andth eViolo-Quercetu m ilicetosum,togethe rwit h Trientalis europaea, Maianthemum bifolium (char acter species for theQuercio nrobori-petraea eaccordin gt oWesthof f& De n Held),an d Pteridium aquilinum. Luzula pilosa, Atrichum undulatum, and Polu- gonatum multiflorum appearedno tt ob edifferentia l forth eViolo-Quercetu m inrelatio n to theQuerco-Betuletum .Thes especie swere ,however ,togethe r with anumbe r ofothe rspecies ,grou p9 i ntabl e1 an d2 ,differentia lfo r theViolo-Quercetu m typicum andilicetosu man dth eQuerco-Betuletu m inre lationt oth eDicrano-Quercetum .Prunu s serotina isno to ra tth emos tver y weaklydifferentia lfo rth eViolo-Quercetu m typicuman dth eQuerco-Betuletu m together inrelatio n toth eDicrano-Quercetu m andth eViolo-Quercetu m ili- • cetosum. Of themosse s growing on dead stumpsan dlogs , Campylopus fragilis ap peared tob e differential forth eViolo-Quercetu m typicum inrelatio nt o theothe r (sub)associations. Dicranella heteromalla, Eurhynchium praelongum, and Plagiothecium latebricola are differential for the Violo-Quercetum typicuman dilicetosu m inrelatio nt oth eDicrano-Quercetu m andth eQuerco - Betuletum. A group of8 species isdifferentia l forth e Violo-Quercetum typicuman dth eQuerco-Betuletu m together: Mnium hornum, Dicranum scoparium, Tetraphis pellucida, Orthodontium lineare, Cladonia glauca, C. digitata, Polytrichum formosum, Plagiothecium curvifolium, and Campylopus flexuosus. Differentialspecie sfo rth eViolo-Quercetu m typicuman dilicetosu man dfo r theQuerco-Betuletu m inrelatio nt oth eDicrano-Quercetu m are Brachythecium rutabulum, Isothecium myosuroides, Lepidozia reptans, and Isopterygium seligeri. 25 Inthi sassociatio nther ewer ecoppice san dhig hwoods ,a si nth eQuerco - Betuletum. Inth ecoppice sth e shrub layerwa sabsent ,o rver yscanty .I n thehig hwood s the shrub layerwa smor e developed, cover15-20% ,tha ni n thehig hwood s ofth e Querco-Betuletum.Th etre elaye ro fth eViolo-Quer - cetumtypicu mdi dno tdiffe rgreatl yfro mtha to fth eQuerco-Betuletum .Th e composition ofth eher b layer differs themos t from thato f theQuerco - Betuletum. Herbs that are aspectdeterminin g inth eQuerco-Betuletu mwil l occuri nth eViolo-Quercetum ,bu tonl yi na spars ecover .Th easpec to fth e herb layer in the Violo-Quercetum typicum is determined by Pteridium aquilinum, Maianthemum bifolium, Trientalis europaea, Oxalis acetosella, terrestrial Hedera helix, Stellaria holostea, and Convallaria majalis. 2.2.3 The Violo-Quercetum ilicetosum Violo-Quercetum ilicetosum subass.nov . type:relev é2 3 (table 1), TheNorgerholt ,prov .o fDrenthe ,Th eNether lands, nonQuerceto-Ilicetu mTx .1930 . TheQuerceto-Ilicetu mTx .193 0i sa completel ydifferen tvegetatio ntype , with not only Maianthemum bifolium and Ilex aquifolium ascharacte ran d differential species,bu tals o Teucrium scorodonium, Thelypteris dryopteris, Luzula luzuloides, and Solidago virgaurea. Herb andshru blayer sar emuc h more developed than those ofth eViolo-Quercetu m ilicetosum, and contain speciessuc ha s Carpinus betulus, Crataegus spec, Prunus spinosa, Athyrium filix-femina, Circaea lutetiana. The only differential species for the Violo-Quercetum ilicetosum in relation toth eViolo-Quercetu m typicumi s Ilex aquifolium inth etre ean d shrublayer .Th eoccurranc eo f Isopterygium seligeri inth eViolo-Quercetu m ilicetosum was called characteristic by Westhoff& De nHel d (1969),bu t appeared not todiffe r from that inth eViolo-Quercetu m typicum andth e Querco-Betuletum,bot h forterrestria lspecimen san dth especimen sgrowin g on dead stumps.Th e Violo-Quercetum ilicetosum and theViolo-Quercetu m typicum have 9differentia l species incommo n inrelatio n toth eQuerco - Betuletum: Oxalis acetosella, Hedera helix, Convallaria majalis, Corylus ave 11ana, Maianthemum bifolium, Trientalis europaea, Stellaria holostea, Pteridium aquilinum, and Ilex aquifolium. Therear en odifferentia lspecie s for theViolo-Quercetu m ilicetosum and theQuerco-Betuletu m incommo ni n relationt oth eViolo-Quercetu mtypicum . 72ex aquifoilium occurred as anearl y closed lowtre e layer orhig h shrub layer.A herb layerwa s almost absent,th emea ncove rwa s7.5 %(i n theViolo-Quercetu m typicum 50%). Inope n spaces and atth e edge aher b layerwa s observed similar to thato fa Violo-Quercetu m typicum.Fo rthi s reason andbecaus e of thenumber s ofdifferentia l speciesI conside rthi s 26 Vegetationt obelon gt oth eViolo-Quercetu m andhav e ranked ita sa sub - association. The almost complete absence ofa her b layer isprobabl y causedb y screeningal lth eyea rroun db y Ilex trees,whic hprevent sth estron gligh t frompenetratin g tothi s level.Th e Ilex vegetations probably originated from aViolo-Quercetu m typicum byspontaneou s growth. This spontaneous growtho fyoun g Ilexshrub swa sals oobserve di nsom eo fth eViolo-Quercetu m typicumplots . TheViolo-Quercetu m ilicetosumi spossibl y restrictedt oth eDrenthia n district (Barkman& Westhof f 1969). Iti s also rather rare withinth e Drenthiandistrict . The investigated plots were allhig hwood swit h large,thu sol doa k trees. 2.2.4 The Dicrano-Quercetum Dicrano-Quercetum Pass.196 2emend .Barkm .n.p . Synonyms: Querco-BetuletumTx .193 0p.p . Vaccinio-Quercetum cladonietosumDoin g1962 . Classification: According toBarkma n (1974)an dt oVreugdenhi l& Barkma n (ReportBiol . Station,Wijster )th eDicrano-Quercetu m inth eNetherland sca nb edivide d intotw osubassociations :plagiothecietosu m Barkm.197 4(wit htw ovariants ) and polypodietosum Barkm. 1974.Th eDicrano-Quercetu m ofPassarg e iss o poori nspecies ,tha ti tca nno tb eplace di non eo fthes esubassociations . In a study onth evegetatio n ofth eDicrano-Quercetu m Vreugdenhil& Barkmanmad e4 2relevé si nth eprovinc eo fDrenthe ,an d1 1relevé si nth e innercoasta ldune snea rAlkmaa r (provinceo fN.-Holland) .Thes e5 3relevé s were compared with relevés onrelate d vegetations, mostly rather poor Querco-Betuletum vegetations, known from literature.The y founda larg e numbero fdifferentia l species,mostl ymosses ,fo rth eDicrano-Quercetu mi n relation toth eQuerco-Betuletum . Icompare dVreugdenhi l& Barkman' s4 2 relevés fromDrenth ewit hth erelevé s givenher eo fth eQuerco-Betuletu m and theViolo-Quercetu m typicum andilicetosum , fromth esam eprovince . There appearedt ob e1 4distinctl y and1 4weakl ydifferentia l speciesfo r the Dicrano-Quercetum in relation toth eQuerco-Betuletum : Dicranum scoparium, Pohlia nutans, Lecidea granulosa, Cladonia puxidata, Parmelia phusodes, Cladonia glauca, Dicranum polysetum, Pleurozium schreberi, Cephaloziella divaricata, Cladonia impexa, Calluna vulgaris, Cephalozia bicuspidata, Cladonia furcata, Cladonia macilenta; respectively Lophocolea heterophglla, Campglopus fragilis, Aulacomnium androgynum, Dicranoweisia cirrhata, Festuca ovina, Empetrumnigrum, Cladonia uncialis, Orthodicranum montaum, Agrostis canina, Dicranum fuscescens, Ptilidium ciliare, Corni- 27 cularia aculeata, Cladonia floerkeana and Cetraria glauca. Allmosse san d lichens named here,wer egrowin gterrestrially .Som eo fthes especie swil l sometimes occur in aQuerco-Betuletu m or ina Violo-Quercetum ,bu tthe n always onbark ,o r dead stumpso r logs. Inth erelevé so fmosse so ndea d stumps and logs,n o species appeared tob e differential forth eDicrano - Quercetumi nrelatio nt oth eothe rassociations . Inthi s studyonl y 3plot s ofth eDicrano-Quercetu mwer einvestigated . Thesynoptica ltabl eo fth eDicrano-Quercetu mwa sno tmad eo fthes e3 rele vés,bu t ofth e4 2relevé smad e inDrenth eb yVreugdenhi l& Barkman .Th e 3relevé s givenher ebelon gt oth e subassociationplagiothecietosum , the relevés 1an d 2belon gt o thevarian to f Cetraria glauca, therelev é3 t o thevarian to f Tetraphis pellucida. The investigated vegetations werescrubb ywoods ,o rsometime scoppices . The tree layer was alwaysver y low,0.5-8(-15 )m high ;withou t ashru b layer.Th eher b layerwa s alwaysver ysparse ,cove r1-12% ,whic hconsite d of Calluna vulgaris, Festuca ovina, Deschampsia flexuosa, and less fre quently of Empetrum nigrum and Carex pilulifera. Thisher blaye rtherefor e lacks the species thatar echaracteristi c forth eQuerco-Betuletu m andth e Violo-Quercetum,an dsom especie soccu rtha tar etypica lo fver ydr yheath s andinlan ddrif tsan dareas .Th emos slaye rwa sver ycharacteristi c forth e Dicrano-Quercetum, thisoccurre d inpatches ,wit h amea ncove ro f70% .I n Drenthe Dicranum scoparium dominated inth emos slayer . 2.2.5 Classification in higher ranks Generally theQuerco-Betuletu m and theViolo-Quercetu m aregroupe d in the alliance Quercionrobori-petraea e (Malcuit1929 )Br.-Bl .1932 ,i nth e orderQuercetali arobori-petraea eTx .1931 ,an dth eclas sQuercete arobori - petraeaeBr.-Bl .e tTx .1934 . Scamoni& Passarg e (1959), Doing (1962, 1963) and Passarge& Hofman n (1968)di dno tagre ewit h thisclassification .Doin gconsidere d thatth e Quercetalia robori-petraeae belonged to the classQuerco-Piceete a Doing 1962, togetherwit hth eBetulo-Vaccinietali a uliginosi and theVaccinio - Piceetalia.Passarg e& Hofman n used theirow nclassification .Th eMolinio - Quercion Scam,e tPass . 1959,th eAgrostido-Quercio nScam ,e tPass .1959 , and theMelampyro-Quercio nPass , etHofm . 1968ma yb econsidere dtogethe r as synonym with theQuercio n robori-petraeae.The y classed theMolinio - Quercion in theorde rMolinio-Quercetali a Pass, etHofm .1968 ;bot hothe r allianceswer e classed inth e orderMelampyro-Quercetali arobori-patraea e Pass, etHofm . 1968.Th e 2order swer eunite dint oth eclas sDeschampsio - Quercetea robori-petraeae Br.-Bl. etTx .194 3em .Pass ,e tHofm .1968 ,to getherwit hth eDicrano-Quercetali a robori-petraeaePass ,e tHofm .1968 . TheDicrano-Quercetu m was classedb yPassarg e& Hofman ni nth eallianc e Dicrano-Quercion Pass. 1963, in the order Dicrano-Quercetalia robori- 28 petraeae, and inth e classDeschampsio-Quercete a robori-petraeae.Barkma n (1974)propose d uniting theDicrano-Quercetu mwit hth eDicrano-Juniperetu m and the Leucobryo-Pinetum to the alliance Dicrano-Pinion. TheDicrano - Quercetumwa sno tmentione db yothe rauthors . Thepositio n ofth eDicrano-Pinio n (Libbert1933 )Matuszkiewic z196 2i s ratheruncertain .Scamon i& Passarg e (1959),Doin gKraf t& Westhof f (1959), andDoin g (1962)unite d theVaccinio-Piceetali a -th e order towhic hth e Dicrano-Pinionshoul dbelon g- togethe rwit hth eQuercetali arobori-petraea e toth eclas sBetulo-Pinete a Prsg.e tKnap p 1942,o r toth eclas sQuerco - Piceetea DoingKraf te tWesthof f 1959.Thi swa srejecte db yMatuszkiewis z (1962)an dWesthof f& De nHel d (1969). Tüxen (1955)an dOberdorfe re tal . (1967)di dno trecognis eth eDicrano-Pinion . Inthi s study a largenumbe r ofspecie swa sfoun dt odifferentiat eth e Dicrano-Quercetum fromth eQuerco-Betuletu m andth eViolo-Quercetum :2 8fo r theDicrano-Quercetu m inrelatio n toth eQuerco-Betuletu m and theViolo - Quercetum,an dtogethe r3 0fo rth eQuerco-Betuletu m andth eViolo-Quercetum , separatelyan dtogether ,i nrelatio nt oth eDicrano-Quercetum .Thes enumber s ofdifferentia l speciesjustif yth eopinio ntha tth eDicrano-Quercetu mdoe s notbelon g to theQuercio nrobori-petraeae .Th especie so fgrou p9 (table s 1 and 2) are to be considered asdifferentia l species ofth eQuercio n robori-petraeaei nrelatio nt oth eDicrano-Quercetum . 2.2.6 Mathematical affinities Thevalue s forA c,A^ ,an d A^, (table3 )an dfo rD c,D p/ andD T (fig.1 ; the length ofth econnection sar emeasure sfo rth edifference sbetwee nth e associations) show that there is a great affinity between theQuerco - Betuletum and theViolo-Quercetu m typicum, evengreate r thanbetwee nth e Violo-Quercetum typicum and theViolo-Quercetu m ilicetosum. Thiswa sun expected as itwa santicipate dtha tth ehighes taffinit yvalue swoul dhav e beenbetwee nbot h thesubassociation so fth eViolo-Quercetum .Th eaffinit y of the Violo-Quercetum ilicetosum was slightly greaterwit h theViolo - Quercetum typicumtha nwit hth eQuerco-Betuletu m inal lcalculations . The affinity of theDicrano-Quercetu m with the othervegetation swa s verylow ,especiall ywit hth eViolo-Quercetu m ilicetosum.Thi swa sexpecte d in relation to the largenumbe r ofdifferentia l species of theDicrano - Quercetum. The affinities ofth evegetation s based onth edifferentia lspecie s( a relative smallnumber )wer ebes testimate db yth evalue sfo rA .Th erela tion of the affinities appeared tochang e if also thePresenc e andth e TotalCove rValu ewer einclude di ncalculatin gmathematica l affinities:th e affinityo fth eViolo-Quercetu m ilicetosumwit hth eViolo-Quercetu mtypicu m orth eQuerco-Betulteu m decreased, and increased ofth eDicrano-Quercetu m withth eViolo-Quercetu mtypicu mo rth eQuerco-Betuletum . 29 Table3 : Mathematical affinities according to3 differen tcalculations . Seetext . *r DQ -QB 0,89 0,62 0,52 DQ -VQt 0,67 0,71 0,46 DQ -VQi 0,52 0,32 0,25 QB -VQt 3,11 3,29 1,29 QB -VQi 1,78 1,71 0,55 VQt-VQi 2,71 2,01 0,62 VQt VQt Fig.1 .Th edifference sbetwee nth evegetatio ntypes ,D ,D p,an dD T,base d onth evegetatio no fplants ,mosses ,an dlichens . 30 2.3 CONCLUSIONS The Querco-Betuletum and theViolo-Quercetu m canb e regarded asver y closely related,bu t independent associations,whic h occur sideb yside . Theybelon gt oth eQuercio nrobori-petraeae .Th eViolo-Quercetu milicetosu m is a subassociation of the Violo-Quercetum because ofth e differential species and inspit e of the low affinity values,lowe rtha nthos eo fth e Violo-Quercetum typicumwit hth eQuerco-Betuletum . The Dicrano-Quercetum has much differential species and low affinity values. Itha s tob e considered asa nindependen tassociatio ntha ti sno t very closely related to the associations ofth eQuercio nrobori-petraeae . Itprobabl ybelong st oth eDicrano-Pinion . 31 3 SOIL 3.1.1 Introduction Soil researchwa s donei norde rt ogiv ea furthe r descriptiono fth e habitato fth ephytocoenoses .Th esoi lprofile san danumbe ro fchemica l factorswer einvestigated . Thesoi lprofil eo fsom eplot swa sknown ,a swel la sth ebi gdifference s between theseprofiles .A mor e detailed surveyseeme dnecessary .Simulta neouslyth egroun dwate rlevel swer estudie dtoo . Thechemica lfactor sanalyse dwer eth epH ,th econductivity ,th eamount s of exchangeable sodium, potassium, calcium, andmagnesiu m ions,an dth e amountso ftota lcarbo nan dnitrogen .Othe rions ,includin gth eanions , are alsoneede dfo rgrowt ho fplants .I nmeasurin gth eamount so fth e4 kations , together with the conductivity,w ehav ea nindication ,bu tno ta complet e description,o fth enutrien tsupply .Fung iar elikel ymor et odepen do nth e amountso fcarbo n andnitroge ntha no nth eamount so fions .Th eamount so f totalC an dtota lN wer etherefor emeasured ,an dth eC/ Nrati odetermined . 3.1.2 A general description of the soil in Drenthe. Froma geologica l pointo fview ,th esoil so fth eprovinc eo fDrenth e arerelativel yyoung ,datin gfro mHolocen ean dPleistocene .Drenth econsist s mainlyo fth eDrenth e Formation, glacial groundmorain e (till)deposits , with some small areaso ffluvioglacia l deposits,datin g fromth eSaal e glaciation (Pleistocene). Iti scovere db ya thi n layero fth eyounge r TwenteFormation .Th eTwent eformation ,datin gfro mth elas tglacia lphase , the Weichselien (Pleistocene),i srepresente db ycoversands ,an di nth e Eastan dSout hEas tpart so fDrenth eb yfluvio-periglacia l peatdeposit s coveredb ya thi nlaye ro fcove rsands .Holocen edeposit sar efoun di nth e SouthEast ,partl yreclaime d depositso fpeat ,o fth eGriendtsvee nForma tion. Scattered throughout theprovinc einlan ddune-sand sar efoun do fth e Kootwijk Formation (Holocene). Inth ebrookvalley sther ear eorgani cbroo k depositso fth eSingarave nFormatio n (Holocene). The influenceo fglacie r activityi sals ob esee ni nth egeomorpholog y ofDrenthe .Th emai npar to fth eprovinc econsist so flo wrise si ngroun d moraine, usuallywit h coversands .Som evalley-lik edepression sar efoun d thathav ebee nmad eb ymeltin gsno wwater ,an ddepression stha tca nb econ sidered- a tleas tpartl y- a spingo-remnants . Some ice-pushed ridgesar e found,usuall y with till,fo rinstanc ei nth eSout hWest ,nea rSteenwijk . 32 The ridges in theEast ,suc ha sth eHondsrug ,wer epossibl y formedb ytec tonicmovements .Th e relief ofDrenth e isals omarke db ylo winlan ddune s (withfla twind-blow nsand yareas )an dhighe rinlan ddunes .I nth eEas tan d SouthEas tther ear eplain stha thav ebee nforme db ypea treclaimation ,an d areaswit helevate dpea tremnants . 3.2 SOILPROFILE S 3.2.1 Methods Soil profiles were taken in april 1978, and again in June 1979.A 120-170c m long and 6c mwid esoilauge rwa sused .Th eprofil ewa sexamine d to adept h of 100-150cm .Th e depth,thickness ,colour ,grainsize ,amoun t ofhumu san dhumu sstructur eo feac hhumu san dminera llaye rwa snoted ,to getherwit hnecessar y particularities.Th elayer swer estudie dmacroscopi - cally in the field, and afterwards with abinocula r loupe (upt o3Ox )i n the laboratory. Colours aredescribe d and coded according to theMunsel l Soil Colour Charts.Th e soilwa s described fresha swel la sdrie dfo rca . 24h a t105°C . Soilswer edetermine d accordingt oD eBakke r& Schellin g (1966). Soilmap sheets 12E (Stiboka 1977), 17E and 17W (Stiboka 1978)scal e1:50.00 0 werestudied . Theleve lo fth egroun dwate ri sgive ni n7 classes ,'grondwatertrappen ' (Gt)o nth e soilmaps, defined after the 'meanhighes tgroun dwate rlevel ' andth e 'meanlowes tgroun dwate rlevel '(Stibok a 1977).Th eG tvalue swer e read from the soilmaps. Ifn o soilmapwa savailable ,th eG twa sestimate d fromth esoi lprofile . 33 Additionalnote so ntabl e4 . Plotnr . 1.A t 85->110c mdepth :a nolde rprofile ,burie dunde rdriftsand . 2.A t110->12 5c mdepth :a nolde rA llayer ,burie dunde rdriftsand . 3.A t 50-10 0c mdepth :a nolde rprofile ,burie dunde rdriftsand . 5.Th eA playe rha dcavitie san dmixe dcolours . 6.Th eA p layerwa s amixtur e of formerA ,B and Clayer s andha dbee n turnedu pwhe nth edrain swer edug. - 7.Th e upper 14c m showed evidence ofhuma nactivit y (digging)an donl ya smallprofil eha sdevelope dove rth eyears .Th eC Ilaye rwa s5 c mthick . 8.Th euppe r 16c mshowe devidenc eo fhuma nactivit yan donl ya smal lpro fileha sdeveloped .Th eA22 ban dth eB2 blayer sha dn oclea rcharacteristics . 9.Th ecove ro fth eA0 0contain si nadditio n8 %Betul apubescens .Ther ewa s also some disturbance onthi splo t asa resul to fdrai ndigging .Th edis turbed layer is,however ,ver ythin .S oi ti sgive na sa nundisturbe dpro file. 10. TheA p layerha dmixe dcolours .Probabl yth ewhol esit ewa sdu gou ta t thetim etha tth edrain swer elaid .I nth euppe r1 5c ma new ,smal lprofil e hasdeveloped . 11.Th e upper 24wa sturne du pwhe nth edrain swer edug .I tha sa clea rA l character. 12. TheA2 1laye r canb e divided intoa 4 c mA21 1layer ,a 1 c mdriftsan d layeran da 5 c mA21 2layer . 23.Th ecove ro fth eA0 0contain si nadditio n25 %Fagu ssylvatica .Ther ei s somevariatio n inthicknes s ofth eA l (±2 cm) , theA2 1 (±2 cm )an dth e A22 (±4 cm )layer . 29.Th eB laye ri slacking .Th eC 2laye rha siro nspot san dgleysymptomes . Annext otabl e4 . Concisedescriptio no fth elayers . A00 Litter:las tyear' sleaves ,twigs ,flower setc .Dat ainclud ethick ness,cove rpercentag ean dproportio no feac htyp eo ftree .I nsom elitera turethi slaye ri scalle dth eL-laye r (Litter). A0 Fermentationlayer :partl ydecompose dlitte rwit hrecognizabl eleaf likestructures .Th ethicknes so fth elaye ri sgiven .Colou rappeare dt ob e veryconstant ,5Y R3/3-2 ,dar kreddis hbrow n(i ndrie dconditio n5 Y E4/ 3- 7.5YR4/2 ,reddis hbrow n to darkbrown) . Insom eliteratur ethi slaye ri s calledth eF-laye r(Fermentation) . Al Humuslayer : decomposed litter, withoutvisibl e plantstructures . Tickness is indicated. Colourappeare dt ob ever yconstan ti nfres hhumus , mostly 5YR2/2 ,sometime s 5YR2/ 1 -3/2-4 ,blac kt odar kreddis hbrow n(i n dried conditionvariatio n issomewha tlarger ,includin g5Y R2/1-2 ,3-4/2- 3 and7.5Y R4/4 ,black ,dar kreddis hbrown ,reddis hbrow nan ddar kbrown) . 34 Sandgrainswer eofte nfoun dscattere di nth eA llayer .Thi slaye ri sclearl y ahumu slayer ,an dno ta minera llaye rwit hinfiltrate dhumus .Th eadditio n ofsandgrain swa s due to animal activity.Th ehumu sappeare dalway st ob e of an amorphous structure,onl y once some 'moder'particle s were found. Humification is apparently caused by fragmentation and growth of fungal mycelia. This type ofhumu s iscalle d 'mor'o r 'rawhumus ' (Wilde1946 , Kubiëna 1953). Insom e soils of theViolo-Quercetu milicetosu m aA llaye r wasmissing , andwa s replaced by aAl hlaye r (seebelow) . Insom elitera turethi slaye ri scalle da H-laye r (Humus). Alh Expeciallyi nsoil so fth eViolo-Quercetu m ilicetosum ablac klaye r was found consisting mainly or entirely of infiltrated, amorphoushumus . Thelaye rwa splasti can dslipper ywhe nwet .Sandgrain swer escars et over y scarse,an dalway sleached .Whe ndrie dver yhar dpiece swer eformed .Colour : black,rarel ydar kreddis hbrown ,5Y R2/1-2 ,7.5Y R2/0 ,10Y R2/ 1 (dryblac k to dark gray, rarely dark reddish brown,5Y R2-4/1 ,2/2 ,10Y R3-4/1) .I have called ita nA llaye rbecaus e of theorgani c originan di nspit eo f the infiltrated character, and added h toindicat eth erichnes si ninfil trated humus.A layero fthi styp eca noccu ro nbadl ydraine dsoil s (Wilde 1946). A21 Mineral layer of leachedsandgrain swit ha fai rt olarg eamoun to f infiltrated humus. The humous part isdominant .Colours :blac k todar k gray, very dark brown, dark reddishbrow n (5YR2/1-2 ,3/1 ,7.5Y R2-3/0 , 10YR2-4/1 ,2/ 2 (dryver ydar kgra yt ogray ,t odar kbrow no rdar kreddis h brown,5Y R3/3 ,5/1-2 ,7.5Y R3-5/0 ,3/2 ,10Y R3-5/1 ,2.5 Y4-5/0-1 ,5/2) . A22 Minerallaye ro fleache dsan dwit hsom einfiltrate dhumus .Th esan d fractioni salway sdominant .Eluvia llaye rwit hpodsoli ccharacter .Colours : gray or grayish brown tover y dark gray orgrayis hbrown ,10Y R3-5/1-2 , 2.5Y4/0 , 3-5/2 (dry dark brown or grayish brown,dar k to lightgray , 10YR3/3 ,4-6/1-2 ,7/1 ,2.5 Y4-7/0 ,5-6/2) . Ap AnA layer ,disturbe db yhuma nactivity ,suc ha sploughing . B Layers characterized by accumulationo fdisperse dhumus ,coatin gth e sandgrains. B2 The layerwit h thebigges tamoun to faccumulate dhumus .Humu srich , blacko rver ydar kgray ,t odar kbrow no rdar kreddis hbrown ,rarel yreddis h yellow, 5YR2/1 ,3/1-3 ,7.5Y R2/0 ,10Y R2-3/1 ,3/ 2 (dryver ydar kgra yt o gray and brown, rarely reddishbrown , 5YR4/1 , 3/2,7.5Y R3-4/0 ,4-5/2 , 10YR3-5/1-2) . B3 A layer with very few, extremely fine humus (microhumus). Dark reddish brown, reddish yellow,yellowis h red,brow n oryellowis hbrown , 5YR3-4/3 , 3/4 -4/6 , 7.5YR4/2- 4 -6/6 , 10YR3/2 ,4/2-4 ,5/4- 6 (drydar k graybrow nt obrown ,o rligh tyellowis hbrown ,7.5Y R4/2 ,5/6 ,10Y R4-5/2-4 , 6/4). BC Atransitiona llayer ,deepe ri tgraduall ybecam elighter . C Moreo rles sunchange dmaterial :coars et ofin esan d (yellowishbrow n 35 to pale yellow), loamy sand, sandy loam, or loam (gray). Sometimeswit h pebbles,ofte nwit hsmal lrusty-brow niro nspots . D Layer below the Claye r of quite differentcharacte r andorigin . Rare,encounte donl yonc e (plot15) . The character ofth e C andth eD laye r isindicate d ontabl e4 :S=sand , LS=Loamysand , SL=sandyloam ,L=gra yloam ,B=re dboulde r claywit hmanga neselenses . Disturbance. Some of the profiles showed evidence ofhuma n activity (ploughing or digging). Iti sno tknow nwhe nth eplot sha dbee ndisturbed , but it was not done recently. Distrubances wereonl y encounted inth e Querco-Betuletum,an dwer eindicate dwit h 'x'i ntabl e4 . The name of the profiles are also indicated: B='Broek'eart h soil, D=micropodzo l in 'Duin'vagu e soil, H='Haar' podzol soil,M='Moerige ' podzolsoil ,V='Veld 'podzo lsoil . 3.2.2 Results The character, thickness,an d deptho f the layers ofeac hprofil ean d synopticalprofile so fth e4 (sub)associationsar egive n (table4) . The soils all developed ina laye r of coversand.Ofte n grey loamwa s found inth e subsoil. Ingenera l thesoil swer elo wo rmediu mhig hlying . Nearly allplot s had adistinc tpodzo lprofile .Th eB horizon twa s dis tinctlydeveloped ;th ehumu swa salway samorphou san dcoate dth esandrains . Thismean s'tha tthes ewer ehumuspodzo lsoils .Fro mheatin gsample so fall e horizonts ofa profil e (anexperimen t carriedou tonl yonce ,o na profil e ofplo t20 )i t appearedtha tth eA2 1laye rha dles siro ntha nth eA llaye r aboveit ,th eA2 2laye rcontaine d no iron,an dth eB 2laye rha dmuc hmor e iron than theA llayer .Apparantl y the iron togetherwit h thehumu swa s washed out and accumulatedi nth eB 2layer .Nevertheless ,iro ncoating so n sandgrainsi nth elaye rjus tbelo wth eB 2wer eno tfoun dhere ,no ri nothe r plots.A n accumulation of ironi n airo npa nwa sno tencounted .S ohydro - morphic characteristics werepresent .N othic ksan do rcla ycove rwa spre sent,th eA llaye rwa sthin ,an d apeat y topsoil ora intermediat epeat y layerwa sgenerall yabsent .A soi lwit hthes echaracter si scalle da 'Veld ' podzolsoi lan dwa sencounte di n1 7plots . Insom eplot s apeat y topsoilwa spresent .Thi smean ti twa sa 'Moer ' podzol soil. In2 plot s a 'Haar'podzo l soilwa s found,a relativel ydr y humuspodzol. Inon eplo t aB laye rwa s absent;thi swa s a 'Broek'eart h soil. Thesoil si nth eDicrano-Quercetu mwer eal lforme di nyoun gdrif tsands . TheC laye r consisted of finet o rather coarse sand;loa mwa sno tfound . Nodistinc tprofile swer efound ,excep tfo rmicropodzo lprofiles ,o fa mea n length of1 3cm .Soil slik ethi sar ecalle d 'Duin'vagu esoils .Olde rpro files,burie dunde ryounge rdrif tsand ,wer eofte npresent .A sth edept ho f 36 these olderprofile swer ever yvariable , alsowithi n oneplot ,the ywer e notmarke d intabl e4 .Th esoil swer ever ydry ,G tvalue sVII . The soils inth eQuerco-Betuletum ,had ,excep tfo rone ,a mor eo rles s disturbed profile.Sometime s the soilsha dbee n dug and anA playe rwa s present. Sometimes drainswer e dug,an d asandcove ro f11-2 4c mthic kwa s dug upbetwee n the drains.Sometime s ashort ,ne wprofil eha dalread yde veloped inth edu gu psoil .Unde rthi ssandcove ra nol dpodzo lprofil ewa s alwayspresent .Bot h 'Veld'podzo lsoil s (6x)an da 'Moer'podzo lsoi l (lx) were observed.Th e onlyundisturbe d profile wasa 'Haar'podzo lsoil .Th e vegetation in this plot was slightly different from that inth eothe r Querco-Betuletum plots. Including the sandcover thatha dbee ndu gup ,th e C layerwa s found ata mea ndept h of 70cm . Ifw eexclud eth esandcover , the Claye rwa s found ata mea ndept h of 60cm ,an dconsiste do ffaintl y fine to rather coarse sand (3x), loam (2x), or intermediates(3x) . The soils inth eQuerco-Betuletu m werewette r thanthos e inth eDicrano - Quercetum:th eG tvalue svarie dfro mV (2x),t oV I (5x)an dVI I (lx). The profiles ofth e soils inth eViolo-Quercetu m typicumwer e always undisturbed andha d easilyrecognizabl ehorizonts .I twer ealway sdistinc t podzol soils,bot h 'Veld'podzo l soils (6x), as 'Moer'podzo l soils (2x) occured,an donc ea 'Haar'podzo lsoil .Th eC laye rwa sfoun da ta naverag e depth of 75cm , andconsiste do fsan d (4x), loamysan dt osand yloa m(5x) , and once redboulde rclay .Compare dwit hth esoil si nth eQuerco-Betuletu m the profiles ofth eViolo-Quercetu m were about 15c m deeper, and lacked greyloa m (withoutsand )i nth eC layer .I twa sremarkabl etha tth e2 plot s with avegetatio n close to theViolo-Quercetu m ilicetosum,als oha dsoil s thatwer ever y similart othos ei nth eViolo-Quercetu m ilicetosum:a rela tive longprofil e anda distinc tAl hlayer .Th esoil si nth eViolo-Querce tum typicumwer e slightly dryer than thosei nth esoil si nth eQuerco-Be tuletum,G tvalue sV (3x),v i (4x),an dVI I (3x)wer efound . The soils inth eViolo-Quercetu m ilicetosumalway sha dundisturbe dpro files.Th eplot sha dpodzo lsoil sexcep tfo ron eplo twher ea 'Broek'eart h soil occurred.Bot h 'Veld'podzo l soils (4x)a s 'Moer'podzo l soils(3x ) werefound .Th eC laye rwa sa ta mea ndept ho f8 2cm ,an dconsiste do fsan d (3x),o rloam ysan do rsand yloa m (5x).Th eprofil eher ewa so nth eaverag e 7c mlonge r than inth eViolo-Quercetu m typicum, and2 2c mlonge rtha ni n theQuerco-Betuletum . TheC laye rwa sloam yo rsand ya si nth eViolo-Quer cetum typicum, and slightly less loamy than inth eQuerco-Betuletum .Al h layeroccurre dmor e often,an dwa s thicker inth eViolo-Quercetu milice tosum than inth eothe r associations.Thi styp eo flaye rca nonl yfor mi n badly drained soils (Wilde 1946). These soilswer eusuall ywette rtha ni n the other associations,G tvalue s III (3x),v (2x),v i (lx),an dVI I (2x) werefound . 37 3.3 SOILCHEMICA LFACTOR S 3.3.1 Methods Samples for analysis ofchemica l factorswer etake ni nseptembe r1978 . Samples were taken ofth eA O and ofth eA2 1layer ,becaus e thehighes t amounto ffunga lmyceliu m andth elarges tmyceliu mgrowt hoccurre di nthes e layers (DeBooi s 1976). Sampleswer etake nfro mth eAl hinstea do fth eA2 1 layeri fth eAl hlaye rwa s1 0c mo rmor ethick ,becaus ei ti sunlikel ytha t adee plyin gA2 1ha sa larg eamoun to fmucelium . The pH-fresh, the pH-H20, thepH-KCl , the conductivity, the Specific Gravity,th eamount so fexchangeabl eNa ,K ,Ca ,an dM gion san dth etota lC andN wer edetermined ,th e 'totalio nsupply 'an dth eC/ Nrati ocalculated . Todetermin e thepH-fresh ,5 sample so fabou t3 0m lsoi lwer etake npe r plot, inbot h layers.Wate rwa sadde dt oa volum eo fabou t4 0ml .On ehou r laterwa s stirred again;i fther ewa san ysan di trapidl ysan kt oth ebot tom.Th ep Hwa smeasure di nth esuspension . To determine the other factors, a more exactmetho dwa sused .Wit h pF-rings 9sample swer e taken of avolum eo f9 9cm 3 eachi neac hplo tan d bothlayers .Th esample so feac hlaye rwer ecombine dint o3 'mixedsamples ' of3 sample s each.The ywer e stored untilthe ywer eanalyse di na freeze r at-25°C .Th enex t stagewa s that theywer e dried at 50°C for5- 7days . After weighing, the specific gravity was calculated. The samples were ground in amill , theA O soils for 10min. , theA2 1soil sfo r5 min .or , ifthe ywer ever ypeaty , also for1 0min .Th een dproduc to fthes etreat mentsi sfurthe rrefere dt oa ssoil . To determine thepH-H 20 and theconductivity ,5 g soi lwa sshake nwit h 50m lwate r (distilled)fo rhal fa nhour .T odetermin epH-KCl ,5 g soi lwa s shakenwit h 50m lI NKC l solution inwate r foron ehour .Measure dwa si n thesupernatant .Al lp Hvalue swer emeasure dt oon ehundredth ;th eaverage s areth e arithmeticmean s ofth ep Hvalues ,an d rounded offt oon etenth . The conductivity valueswer ecorrecte d forth epH ,s olessene db yth econ ductivityo fth eH ions.Conductivit yi sexpresse di npS/cm . To determine the amounto fexchangeabl eNa ,K ,M gan dC aions ,6 g soi l wasshake nwit h4 0m lNH 4OAc (ammoniumacetaat )p H7.0 ;washe dsuccessivel y with 16,12 ,an d 12m lNH 4OAc over aSchleiche r& Schui ln o58 9blackban d filter, filtred through aSartoriu s 0.8p mmembran efilter ,an dfille du p withNEjOA c solution to 100ml .Thes e solutionswer e used,usuall yundi luted occasionally diluted once,t odetermin e theN aan dK io nconcentra tion in a flame-photometer.Fo rdeterminatio no fth eM gan dC aio nconcen trationi n aPerkin-Elme r atomic absorption spectrophotometer,a dilutio n wasmade : 1m lextrac to fA 0soil ,an d5 m lextrac to fA2 1soil ,wa sadde d to 5m lLa(N0 3)3 solution in water (containing 10,000m gL ape r1 )an d filled upwit hwate rt o5 0ml .Th eamoun to fexchangeabl eion si sgive ni n 38 massfractio n (mg/100g soil )an di nmas sconcentratio n (mg/1soil) . To determine the amounto fcarbo nsom esoi lwa sdrie dagai na t105°C .A portiono fsom egram so fthi ssoi lwa sweighe dan dth elos so nignitio nwa s measuredafte rheatin gt o800° Cfo r 2%hour .Th eamoun to fcarbo ni s58 %o f theweigh tlos s (VanBemmelen-factor) .Th etabl egive sth eamoun to fcarbo n inmas sfractio n (g/100g soil )an di nmas sconcentratio n (g/1soil) . To determine the amounto ftota lnitrogen ,som esoi lwa sdrie dagai na t 105°C. Soil ofA 0 (0.200g )an d ofA2 1 (0.500g or1.00 0 g)wa sweighed , and heated in an acid mixture of6 g salicylic acid in 110m l diluted sulphuricaci d (90m lwate ro n50 0m lH 2S04 s.g.1.84 )a t400° Cfo r1 hour . Thesolutio nwa schille dan ddilute dwit h7 5m lwater .I na Tecato rKjelte c System100 2Distillin gUni t5 0m lNaO H4 %wa sadded .Thi ssolutio nwa sdis tilled into 25m lbori c acid 4%wit hmixe d indicators,an dtitrate dwit h KH (I03)2 0.01N .Th eamoun to fnitroge ni sgive ni nmas sfractio n (g/100g soil)an di nmas sconcentratio n (g/1soil) . 3.3.2 Results The average of3 observation si sgive npe rplo tan dpe rfacto r (forth e pH-fresh the average of5 observations )(tabl e 5).Pe r (sub)associatiepe r factor the average of all observations isgiven ;th eextrem evalue s are mentionedi nparentheses . 3.3.2.1 pH-freshan dpH-H 20 The soilswer e allver y acid.Th ep Hvarie d from3. 0 to4. 7 inth eA 0 layer, and from2. 8 to4. 4 inth eA2 1layer .Ther ewa slittl edifferenc e between the associations.Th evalue s inth epH-fres hvarie dmor etha ni n the pH-H20, because more samples weremeasure d and themetho d was less exact. Inth eA2 1laye rth evalue so fth epH-H 20wer ealway shighe r(aver age 0.6) than those ofth epH-fesh .Als o inth eA 0laye r therewa ssom e differencebetwee nth epH-H 20an d thepH-fres hwithi non eplot ,bu tther e wasmor evariation ;bot hlowe ran dhighe rvalue soccurred .Thes edifference s wereprobabl y causedb y the degree ofdilution .I nmeasurin gth epH-fres h the soilwa s only diluted slightly andwa s measured in asuspension .I n measuring thepH-H 20 the soilwa s dilutedmore .I nth esample so fth eA2 1 layer,whic hwer emor e or less sandy, adistinc t supernatantwa sformed ; the measurements were made in the supernatant. Inth eA 0layer ,usuall y peatyan dno tsandy ,ther ewa salmos tn odeposi tan dth esoi lremaine dmor e orles si nsuspension ;th emeasurement sha dt ob etake ni nte hsuspension . 3.3.2.2 pH-KCl ThepH-KC lvarie d from 2.5 to3. 3 inth eA 0layer .I twa salway slowe r 39 thanth epH-H 20,mea ndifferenc e0.8 .Ther ewa sno tmuc hdifferenc ebetwee n the associations.Th epH-KC lvarie d from 2.4 to 3.5 inth eA2 1layer .I t wasalway slowe rtha nth epH-H 20,mea ndifferenc e0.9 .I nth eseri eDicrano - Quercetum, Querco-Betuletum, Violo-Quercetum typicum, Violo-Quercetum ilicetosum therewer e decreasing pH-KClvalues .Th edifference s wereno t very large,0.5 ,bu t itca nb econclude dtha tth eA2 1laye ri nth eViolo - Quercetum ilicetosumwer eslightl ymor eaci dtha ni nth eDicrano-Quercetum . Theywere ,however ,o fquie ta differen tcharacter :i nth eDicrano-Querce tum itconsiste dmainl y of sand, inth eViolo-Quercetu m ilicetosummainl y ofinfiltrate dhumus . Therelatio npH-KC l andpH-H 20i snearl ylinea ri nbot hth eA 0an dth e A21laye r (fig.2) .Th evalue s ofon e seriep Hmeasurement s arenormall y distributed, itwa s thuscorrec tt otak eth e arithmetical meano fth ep H valuesi ncalculatin gth emea np Hvalues . pH-KCl 3.5- • • • • • e 3.0- o» eft • CD D * a a o e a» 2.5- «e a DQ • QB o VQt a VQi 3.5 LS pH-HjO Fig.2 .Th erelatio nbetwee npH-H 20an dpH-KC li nth eA2 1layer . 40 3.3.2.3 Conductivilyan d 'total'io nsuppl y The conductivity wasrathe rhig h inth eA 0laye rcompare dwit htha ti n theA2 1layer .Th emea nconductivit yvalue si nth eA 0laye rwer ehighe ri n theDicrano-Quercetu m and inth eViolo-Quercetu milicetosu mtha ni nViolo - Quercetum typicum and inQuerco-Betuletum .Th edifference swer elarg ean d werecontrar yt oinitia lexpectation stha tth econductivit ywoul db eequall y highi nal lassociations ,o rlowe ri nth eDicrano-Quercetu m andth eQuerco - Betuletum than in theViolo-Quercetum . Although there is aconsiderabl e variationbetwee n the observations,ther e is asignifican tdifferenc ebe tweenth emean so fth esevera l (sub)associations(F-tes to nlog-transforme d observations:F=4.14 ;3 ,2 5df ;P > 0.025) . The conductivity was relatively low inth eA2 1layer . Inth eDicrano - Quercetum theconductivit y wasobviousl yles stha ni nth eQuerco-Betuletu m and in the Violo-Quercetum typicum, and the conductivity inth eViolo - Quercetum ilicetosum was obviouslyhighe rtha ni nth eQuerco-Betuletu man d inth eViolo-Quercetu mtypicum .Th edifference sbetwee nmean sar estatisti cally highly significant (F-testo nlog-transforme d observations:F=12.7 ; 3,2 5 df;P < <0.005) . There isa distinc tpositiv e correlationwit hth e amounto fcarbo n (fig. 3). The conductivity is ameasur e for the amounto f available ions ina given soilvolume .Th etota l ion supply depends also onth ethicknes so f the layer.T o estimate the 'total'io n supplyth econductivit yo fth eA 0 layerwa smultiplie d by the thickness ofbot hth eA 0an dth eA llaye rto gether (ionsuppl y inA0+A1) ,th econductivit yo fth eA2 1laye rwa smulti pliedb yth ethicknes so fbot hth eAl han dth eA2 1laue rtogethe r (ionsup plyi nAlh+A2) .Th esu mo fbot hvalue si sth e 'total'io nsuppl y (table6) . Inthi s estimation Iassume d that theconductivit y inth eA llaye rwa sa s largea si nth eA 0layer ,an di nth eAl hlaye ra slarg ea si nth eA2 1layer . Iti smor e likely thatth e conductivity inth eA llaye ri slowe rtha ni n theA 0layer ,s o thismultiplication s gives ato ohig hvalue .Th econduc tivity inth eA2 1laye ri s supposed tob elowe rtha ni nth eAl hlayer .I n mostplot s anAl hlaye rwa s notpèsent , so thismultiplicatio nwil lgiv e the accurate value. Ifa thi nAl hlaye rwa s present,th evalu e willb e slightly toolow ;i fa thic kAl hlaye rwa spresen tan dth econductivit yo f thislaye rwa sestablished ,th evalu ewill "b eslightl yto ohigh . The ion supply in the A0+A1laye r is aboutequall y inth eDicrano - Quercetum, the Querco-Betuletum and theViolo-Quercetu m typicum, andi s much higher in the Violo-Quercetum ilicetosum. The ion supply in the Alh+A21laye r is very low in theDicrano-Quercetum , muchhighe r inth e Querco-Betuletum and inth eViolo-Quercetu m typicum, and stillhighe ri n theViolo-Quercetu m ilicetosum. The 'total' ion supply ishighes t inth e Violo-Quercetum ilicetosum, is lessb yhal fi nth eViolo-Quercetu mtypicu m and theQuerco-Betuletu m thatalmos tequal seac hother ,an di sstil llowe r 41 Table6 : 'Total' ion supply. Ion supply in the A0+A1layer , in the Alh+A21layer , and the 'total'io n supply, per plot and per vegetationtype . VQi 29 24 32 56 28 3 6 9 27 60 92 152 26 22 3 26 25 49 31 80 24 2 19 21 23 26 2 28 22 35 11 46 VQt 21 12 18 30 20 9 7 16 19 10 3 13 18 10 2 12 17 11 2 13 16 15 1 16 15 13 6 20 14 20 1 21 13 39 7 46 12 12 2 15 QB 11 7 6 13 10 18 3 21 9 10 10 20 8 20 0.7 20 7 9 2 12 6 31 11 42 5 26 21 47 4 7 3 10 DQ 3 9 0.5 9 2 11 0.1 11 1 23 0.9 24 VQi 28 25 52 VQt 15 5 20 QB 16 7 23 DQ 14 0. 5 14 M 0) u >i i OliH 4> IS as: ID i-H iH S iH H iH rH +< a O +< CM iH 0. conductivity in yuS 200- 1S0- SO' a a • 00 • QB a VQt a VQi •#.•« • SA- 10 IS Carbon(mas s concentration) Fig.3 .Th erelatio nbetwee nth econductivit y andth eproportio no fcarbo n inth eA2 1layer . inth eDicrano-Quercetum .A distinctserie s from a larget oa small ion supplyca nb edistinguished . 3.3.2.4 Specificgravit y The specific gravity inth eA 0laye r isver y constant inal ltype so f vegetation. Inth eA2 1laye r thevariatio n inth e Sg iseasil yexplaine d fromth eamoun to fhumus , (fig. 4). 43 Specific gravity ISA m DQ • QB e VQt o VQi 15 20 Carbon (mass concentration) Fig. 4.Th erelatio nbetwee nth eSpecifi cGravit yan dth eproportio no f carboni nth eA2 1layer . 3.3.2.5 Na,K ,Mg ,an dC Aion s The amounts of available ions isofte ngive ni nmas sfractio n (mgion / 100g soil) . Itseem smor e likely thatth eroot so fplant so rth emyceli a offung ihav ea certai nvolum ei nth esoi lan dar enot ,o rlittle ,affecte d byth eweigh to fth esoil .Becaus eo fthi si ti sbette rt ogiv eth eamount s inmas sconcentratio n (mgion/ 1soil) . Inth eA 0laye r the amount ofN a and ofK ion swa sslightl ylarge ri n theViolo-Quercetu milicetosu mtha ni nth eothe rassociations ,wher ei twa s almost the same.C a andM g ionswer eavailabl ei nabou tth esam equantit y inal l(sub)associations . Inth eA2 1laye r theamoun to fN aion swa slarger ,an dth eamoun to fK , Mg, and Caion swa s smaller,compare d withth eA 0layer . Ingenera lth e variation in amount of ions inthi slaye rwa slarge ,an dther ewa sa con siderableoverla pbetwee nth evalue si nth e (sub)associations.Nevertheles s some differences existed. The amount ofN aion swa slarges ti nth eViolo - Quercetum ilicetosum, andwa s almost the same inth e otherassociations . 44 The amount of Kion s was largesti n theViolo-Quercetu m ilicetosvunan d decreased inth e serieQuerco-Betuletum ,Violo-Quercetu m typicum,Dicrano - Quercetum. The amount ofM gion s decreased inth e serieViolo-Quercetu m ilicetosum, Querco-Betuletum, Violo-Quercetum typicum, Dicrano-Quercetum; itwa s distinctly smalleri nth eDicrano-Quercetu m thani nth eothe rasso ciations.Th e amount ofC aion swa s largesti nth eViolo-Quercetu milice tosum, and smaller inth eViolo-Quercetu m typicuman dth eQuerco-Betuletu m -wher e itwa s almost the same- , andwa sconsiderabl y smalleri nth eDi crano-Quercetum. TheViolo-Quercetu m ilicetosum generallyha dth elarges tamount so fions . Thosei nth eViolo-Quercetu m typicuman di nth eQuerco-Betuletu mwer elower , andha dabou tth esam evalues .Thos ei nth eDicrano-Quercetu mwer egenerall y lowertha ni nth eQuerco-Betuletu m andth eViolo-Quercetu mtypicum . A linear correlation with high correlation coefficients appearedt o existbetwee n the ions (inmas s fraction)an d carbon (inmas s fraction). Thecorrelatio no fion san dcarbo nexpresse di nmas sconcentratio nwa sals o linear,bu t the correlation coefficients were slightly lower (fig.5 an d 6). Iti stherefor epossibl et ous eth eamount so fcarbo na sa nestimat eo f theamount so fions . Na-ions (massfraction ) • OQ • QB a VQt a VQi ,• «,«o*/ • 30 40 Carbon(mas s fraction) Fig.5 .Th erelatio nbetwee nth eamoun to fN aion san dcarbo ni nth e A21layer . 45 Na- ions (mass concentration) • DQ • QB o VQt o VQi <. 10 15 Carbon (mass concentration) Fig.6 .Th erelatio nbetwee nth eamoun to fN aion san dcarbo ni nth e A21layer . 46 a cl o2o >2 >o o oo CT«I Z° Fig.7 .Th erelatio nbetwee nth eproportion so fnitroge nan dcarbo ni nth e A21 layer (left) and in the AO layer (right). 47 3.3.2.6 Totalcarbo nan dnitroge n Inth eA Olaye rth eamount so fcarbo nan dnitroge nbot hdecrease di nth e serieViolo-Quercetu milicetosum ,Violo-Quercetu mtypicum ,Querco-Betuletum , Dicrano-Quercetum, but the differences were rather small.Als o the C/N ratiodiffere dver ylittle ,i tincrease dslightl yi nth esam eserie . Inth eA2 1laye r the difference between theassociation swa smor edis tinct.Bot hth eamoun to fcarbo nan dnitroge nwa slarges ti nth eViolo-Quer cetum ilicetosum, smaller and aboutequa l inth eViolo-Quercetu m typicum andth eQuerco-Betuletum , and smallesti nth eDicrano-Quercetum . The C/N ratiowa shighes t inth eDicrano-Quercetu m andth eQuerco-Betuletum ,wher e itwa s almostequal .I twa slowe ran dals oalmos tequa li nth eViolo-Quer cetum ilicetosum. There isa stron glinea rcorrelatio nbetwee nth eamount s of nitrogen and carbon, inbot hth e layers (fig.7) ,comparabl e toth e correlationo fth eamount so fion san dcarbon . 3.4 CONCLUSIONS The soils inth efou r (sub)associations differgreatl yfro meac hother . Differences were found inth e soilprofile s and inth echemica lfactors , especially inth eA2 1layer . Inth eViolo-Quercetu m ilicetosum theplot s had relatively deepprofile s with athic kAl hlayer ,th ehighes tio nsup plies,th elowes tpH ,th ehighes tNa ,K ,Mg ,an dC aio nconcentrations ,th e highestproportion s ofnitroge nan dcarbon .Th eC/ Nratio' sar erelativel y low and equal thosei nth eViolo-Quercetu m typicum. Inth eViolo-Quercetu m typicum and the Querco-Betuletum theplot s hadmor e shallowprofiles ,i n theQuerco-Betuletu m slightly moreshallo wcompare dt oth eViolo-Quercetu m typicum.Th etopsoi l inth eQuerco-Betuletu mplot sgenerall ywa sdisturbe d by formerhuma n activity.Th epH ,th eK an dth eM gio nconcentrations ,an d the C/Nratio' s were slightly higher,th e Caconcentratio nwa s slightly lower inth eQuerco-Betuletu m comparedt oth eViolo-Quercetu m typicum,th e ionsupply , theN a concentration, theproportion s ofnitroge n andcarbo n werealmos tequa li nbot hth evegetatio ntypes .I nth eDicano-Quercetu mth e plotsha dver y shallowprofiles ,guiet e differentfro mthos ei nth eothe r associations.Th ep Hwa shigher ,th e ionsupply ,th econcentration so fK , Mg,an dC aions ,an dth eproportion so fnitroge nan dcarbo nwer elowe rtha n inth eothe r associations.Th eN a ionconcentratio n is almostequa lher e andi nth eViolo-Quercetu m typicum andth eQuerco-Betuletum .Th eC/ Nratio' s wererelativel yhig han dequa lthos ei nth eQuerco-Betuletum . 48 THE MYCOCOENOSES 4.1 INTRODUCTION To describe the fungusvegetatio no fphytocoenose ,i ti sno tsufficien t tomak ejus ton erelevé ,jus tlik eth edescriptio no fa phytocoenos eb yon e relevé of thephaneroga m vegetation.Fung i havemuc hmor epronounce dsea sonalaspect stha ngree nplants :th elat esumme raspec tdiffer spartl yfro m the early autumn aspectan dt oa larg eexten tfro mth elat eautum naspect . A fungusvegetatio nma yals osho wbi gdifferences ,qualitativel yan dquan titatively, from year toyear .Thi s depends agrea t dealo nth eweather , especially therain ,o nhumidit y and temperatureo fai ran dsoi l(Barkma n 1976b,Thoe n 1976), onth eweathe ro fprecedin gperiods ,an dpossibl yals o onother ,partl yunknow nfactors . 4.2 METHODS 4.2.1 Relevés and tables To get acquainted with the whole fungusvegetatio n of aplo t iti s necessary tomak e anumbe r ofrelevé seac hyea rfo rsevera lyears .I twa s planned tomak e 2o r3 relevé spe r fructification season (latesumme ran d autumn)i neac hplo tfo r4 years .Thi sgoa lwa sno talway sachieved ,partl y because the frequency relevés,an dsometime sdetermination swer ever ytim e consuming,partl ybecaus e of fluctuations in fructification (forinstanc e in197 6fructificatio ndi dno tstar tunti lth ebeginnin go fOctober) . Theplot swer eno tvisite dofte ni nth eothe rseasons .I nwinter ,spring , and early summer thenumbe r of fungiwa salway sver ylow ,mostl yver yol d specimens from the preceding autumn. Observations on fungi inwinter , spring, and early summerwer e alsouse d asrelevé s andworke d intoth e tables. In a relevé, abundance was determined by counting orestimatin g the number ofcarpophore s ofeac h fungus species (abundancerelevé) .Countin g thenumbe r ofcarpophore s is amor eexact ,bu tmor etim econsumin gmetho d thanestimating .Her e acombinatio n ofbot hwa sused .Abundanc ewa snote d in the following scale (according to Barkman 1976, slightly modified; mentionned inparenthesi s areth emea nnumbe ro fcarpophore so ftha tclass) : r (rare) 1o r 2carpophore s (1.4) o (occasional) 3- 9carpophore s (5) f (frequent) 10- 29carpophore s (17) 49 vf (veryfrequent ) 30- 99carpophore s (54) a (abundant) 100- 50 0carpophore s (223) va (veryabundant )mor etha n50 0carpophore s The abundancewa s countedi na nare ao f100 0m 2. Inplot stha twer elarge r orsmalle rtha n100 0m 2, abundancewa sconverte dan dals ocounte di nnumbe r ofcarpophore spe r100 0m z. Thenumbe r ofcarpophore swa sno tdetermine d forsom especie swit hver y small fruitingbodies .Her e thenumbe r of (small)growin gspot swa snote d instead. This was never applied to Agaricales, only to speciesgrowin g clustered onwoo d or anothe r limited substrate,suc h as species ofth e genera Hymenoscx/phus, Cudoniella, Sphaerobolus, Onygena, Ciboria, andt o species,growin g onwood ,whos e carpophores areno tseperat eindividuals , sucha sspecie so fth egener a Stereum, Phlebia, Merulius, Exidia. Anabundanc etabl ewa smad eo feac hplo twher eth especie san dth eabun dancewer e giveno feac hrelevé .Dat awer e also included ofearlie rre search on theseplot s (Barkmanpers . comm.,Ypelaa r inprep.) . Inthes e tables itwa s easyt orea dth eMaximu mAbundanc eo fCarpophore s (MAC)o fa speciespe ryear ,an dth eAbsolut eMaximu mAbundanc eo fCarpophore s (AMAC), thehighes t ofal lMA Cvalue s oftha tspecie s (Barkman1976) . Itwa sals o easyt orea dth enumbe ro fyear si nwhic ha specie swa sfound ,s ot odeter mine itsAnnua l Frequency (AF) (Barkman 1976).Value sfo rAMA Can dA Far e giveni ntabl e7 . In1 7plot sno tonl yth eabundanc eo fth efungu sspecie swa sdetermined , but also the SpatialFrequency .Thes eplot s- frequenc yplot so rcomposit e plots- wer e forthi spurpos e divided intopartia lplot so f2 5m 2.S oth e composite plots of 1000m 2 were divided in 40partia l plots,thos e of 1050m 2 in4 2partia lplots ,an d the oneo f87 5m 2 in3 5partia lplots .A frequency relevé consisted of35 ,40 ,o r4 2partia l relevés,on eo feac h partialplot .Pe r frequency relevé iti seas yt orea di nho wman ypartia l relevés aspecie swa s found,s ot odetermin e its Spatial Frequency:th e proportion (inpercentage )o f the areao fa composit eplo twher eth espe cieswa sfoun di non erelevé . The1 7composit eplot swer edistribute dove rth edifferen tassociations : 2i nth eDicrano-Quercetum ,5 i nth eQuerco-Betuletum ,7 i nth eViolo-Quer - cetum typicum, and 3i nth eViolo-Quercetu milicetosum .Ther ewer ecompar atively few frequency relevésmad ea si trequire da grea tdea lo ftim ean d work: 6i n theDicrano-Quercetu m (3pe rcomposit eplot) , 19i nth eQuerco - Betuletum (3.8pe r compositeplot) ,2 3i nth eViolo-Quercetu m typicum (3.3 percomposit eplot) , and1 0i nth eViolo-Quercetu m ilicetosum (3.3pe rcom positeplot) .Th efrequenc yrelevé swer emad ei nth elat esumme ran dautum n of197 6 (12frequenc yrelevés) ,o f197 7(28) ,an do f197 8(18) . As the partial plots were fixed (with pickets)an dnumbere d itwa s possible to combine the frequency relevés of acomposit eplo t intoon e total frequency relevé. Inthi swa y itwa spossibl et odetermin ei nwhic h 50 andho wman yo fth epartia lplot sa specie swa sfoun ddurin gthes e3 years . I have called the total number ofpartia lplot swher ea specie swa sfoun d theTota l Spatial Frequency. TheTS Fvalue so fth especie sar egive ni na proportion (percentage)o f the areao fth ecomposit eplo t (table 7). The TSFgive sth eproportio no fth eplo tare awer eth ecarpophore so fa specie s werefoun dan di sa nestimatio no fth ecove ro fth emycelia . (seep .68) . Ina frequency relevé thenumbe r ofcarpophore s was alsocounte d (for 1-3 carpophores) or estimated (for4 carpophore s andmore , in 5classes , seep .76 )o f each species ineac hpartia lplot .S oth eabundanc eo fth e species was determined too,an d included inth e abundance table oftha t plot. Some synoptical characters were calculated for each (sub)association (table 8): thePresence ,P ;th eMea nAbsolut eMaximu mAbundanc eo fCarpo phores,GAMA C (Barkman1976) ;th eMea nAnnua lFrequency ,MA F (Barkman1976) ; andth eMea nTota lSpatia lFrequency ,MTSF . To calculate GAMAC, the AMAC valueswer e first reconverted intoth e number ofcarpophores ,usin g thevalue s mentioned inth e abundancescal e (p.48 )o rth erea lnumbe ro fcarpophore s (ifthe ywer ecounted) .Th eGAMA C was calculated over allplots ,includin gth eplot swher ea specie sdi dno t occur, sowher e itsAMA C was 0.Th e GAMACvalue s wereno t given inth e abundance scale,bu t inth enumbe ro fcarpophore spe r1.00 0m 2 roundedof f towhol e figures (ifsmalle rtha none ,rounde dof ft oon etenth) . TheMA Fi sth emea no fth eA Fvalue so fa specie si na (sub)association . TheA Fwa s given inproportio n ofyear s (percentage).Th eMA Fi sth emea n proportion ofyear s (meanpercentage )i nwhic h aspecie swa s found.Als o theMA Fwa s calculatedove ral lplots ,includin gth eplot swher ea specie s didno toccur ,s owher eit sA Fwa s0 . TheMTS F isth emea no f theTS Fvalue so fa specie si na (sub)associa tion.Th eTS Fwa sgive ni nproportio no fth eplo tare a (percentage).S oth e MTSF is the meanproportio n ofth eplo t area (meanpercentage )wher ea specieswa s observed. TheMTS Fwa sonl ycalculate do nfrequenc yplots ,bu t includingplot swher ea specie sdi dno toccur ,s owher eit sTS Fwa s0 . ThePresence ,P , another synoptical chatacter indicatesth eproportio n ofth eplot swher ea specie swa sfound .I ti sth esam echaracte ra suse di n thesynoptica ltabl eo fgree nplants .Presenc ewa scalculate d inpercentage . Iti sgive ni na scal eo f1 0part si nth eRoma nnumeral sI - X . Inth eDicrano-Quercetu mth esynoptica ltabl ewa scalculate do n1 1plots : onth eplot s1 ,2 ,an d3 o fthi ssurve yan do n8 othe rplot sinvestige db y Ypelaar.I nth eothe r (sub)associationsth esynoptica ltabl ewa scalculate d only on theplot sreporte dhere .Th edat ao fearlie rresearc ho nth eplot s byBarkma nusuall yconcerne donl yth epresenc eo fspecie san dno tabundanc e andwer e only included inth ecalculatio no fP ,no ti nth ecalculatio no f theothe rsynoptica lcharacters . The formulas used tocalculat e themathematica l affinities (p.12 )ca n 51 also be used tocalculat e the affinities ofth e fungusvegetations .Th e formulas forth eA andth eA _wer euse dunchanged ,fo rth eA _no tth eTC V values wereused ,bu tth eGAMA Cvalues .Th edifference sbetwee nth efungu s vegetationso fth evegetatio ntype sD ,D p,an dD _wer eals ocalculated . 4.2.2 The increase in the number of species per year of investigation Notal lfungu sspecie swer efoun dever yyear ,an deac hyea rspecie swer e found thatwer eno tpreviousl yobserved .Th etota lnumbe ro fspecie sfoun d ina plo tincrease dwit heac hyea ro finvestigation . Ineac hplo tth enumbe r ofspecie swa s determined afterone ,afte r2 , andafte r3 year so fresearc ha sa proportio n (percentage)o fth enumbe ro f species observed after4 year s ofresearch .Th emean so fthes evalue spe r associationwer e also calculated (Fig.8) .Dat a fromearlie rresearc han d the incomplete data fromth eplot s6 ,7 ,27 ,an d2 9 (theseplot swer eonl y visited for3 years )wer e omitted. Inth eDicrano-Quercetu m the increase was also calculated includingth eobservation smad eb yYpelaa ri n197 2an d 1973. In every followingyea rth e increase inth enumbe r ofspecie sbecam e less, except ina yea rver y rich inspecies .Th e increase inth efourt h year varied from 7t o 10%,an dwa smuc h lowertha nth e increase inth e thirdyea ro f1 0t o24% .I nth eDicrano-Quercetu m theincreas econtinue dt o decrease inth e fifth and sixthyear ,t o4 %i nth e fifthan d 6% inth e sixthyear .Th e sixthyea rwa sapparentl ya year -richi nfungu sspecie si n theDicrano-Quercetum .Barkma n (SyllabusUniv .o fUtrecht )foun da naverag e increaseo f13 %i nth efourt hyea ro nplot si nth eDicrano-Juniperetu mstu died during 13years . Ifoun d asmalle r increase afterth e fourthyear , average8.5% ,probabl ybecaus e Ivisite dth eplot ssevera ltime sa year . Barkmanha d after4 year sonl y found 54%o fth enumbe ro fspecie stha t werepresen ti nth eDicrano-Juniperetu m after1 3year sresearch .Thi smean s that aninvestigatio n lasting only4 year s isno tenoug ht ofin dal lth e fungi in aphytocoenose .Probabl y Ifoun da large rproportio no fth espe ciesi nthes e oakwoods,becaus e Ivisite dth eplot ssevera ltime sa year . Iti sprobabl y comparable with the 80%foun db y Barkman in theDicrano - Juniperetum inth e yeari nwhic hth eincreas ei nth enumbe ro fspecie swa s 9%. Ifw e extrapolate the curves of fig.8 t oa nasymptoti cen dvalu ethi s alsoyield s about80 %o fth een dvalu eafte rth efourt hyear .I ti sthere fore notunlikel y that the tables represent about80 %o fth etota lmyco - florao fth eplot sinvestigated . 52 o —t CM O UI I O ». — .2 M ?3= 03 O5 . E -*3 Fig.8 .Th emea nnumbe r of species inth e (sub)associationsafte r1 ,2 ,3 and 4year s ofresearc h as aproportio n (percentage)o fth etota l number of species.1=1976 ,2=1976+1977 ,3=1976+1977+1978 ,4=1976 + 1977+1978+1979.Right :ide m inth eDicrano-Quercetu mincludin gth e data from earlier research.1=1972 ,2=1972+1973 ,3=1972+1973+1976 , 4=1972+1973+1976+1977, 5=1972+1973+1976+1977+1978, 6=1972+1973+ 1976+1977+1978+1979. 53 Annext otable s7 an d8 . Speciestha twer efoun donl yonce .Mentionne dafte rth enam ear eth eAMAC , theAF ,an d- sometime s- th eTSF .Mentionne di nparenthese sar eth eGAMAC , theMAF ,an d- somtime s- th eMTSF .Th ePresenc eo fthes etax ai salway sI . Plot1 : Cantharellus cibarius var. amethysteus r1 7(0.1 ,1) , Clitocybe cerrusata o1 7(0.4 ,1) , Psathyrella cotonea r1 7(0.2 ,) .Plo t2 : Corti- narius hinnuleus o17, 8(0.7 ,1 ,4) , Galerina cf cerina r17, 3(0.1 ,1 , 1), Inocybe margaritispora o1 7(0.4 , 1), Tricholoa aes tuans r1 7(0.1 , 1), Cortinarius pseudosalor r1 7(0.1 ,1) . Plot3 : Cortiaarius pluvius r1 7 (0.1, 1), Inocybe lanuginella r1 7(0.1 ,1) ,Russula mairei r1 7(0.1 , 1), R. raoultii r1 7 (0.1,1) , R. laurocerasi r1 7 (0.2,1) , Cortinarius hemitrichus r1 7(0.1 ,1) ,Panaeolus fimicola r17 ,3 (0,1 ,1 ,1) , Coltricia perennis r1 7(0.1 ,1) , Sarcodon underwoodii r1 7(0.1 ,1) , Hydnellum conpactum r1 7(0.1 ,1) , Phellodon confluens r1 7(0.1 ,1) ,P. niger r1 7 (0.1, 1).Plo t4 : Amanita porphyria r25 ,1 (0.1 ,3 ,0.2) ,Clitocybe odora o25 ,3 (0.4 ,3 ,0.6) ,Dermocybe crocea o25 ,1 (0.6 ,3 ,0.2) , Hebeloma crustuliniforme r25 ,3 (0.1,3 ,0.6) , Psathyrella gossypina o25 ,3 (0.8, 3,0.6) , P. fusca r25 , 3(0.1 ,3 ,0.6) ,Entoloma farinogustus o25 ,3 (0.4,3 ,0.6) ,Hebeloma velutipes r25 ,3 (0.1 ,3 ,0.6) .Plo t5 : Cortinarius cf fasciatus r25 ,2 (0.1,3 ,0.4) .Plo t6 : Clitocybe angustissima r2 5 (0.1,3) ,Galerina inversa o2 5(0.6 ,3) .Plo t7 : Clitocybe lohjaënsis o3 3 (0.4,4) ,Russula brunneoviolacea 2-X-1961leg .J.J .Barkma n708 4(Herb . WAG-W). Plot8 : Cantharellus sinuosus f2 5(3 ,3) , Galerina cf incurvata r2 5(0.2 ,3) , Cyathipodia macropus o2 5(0.4 ,3) ,Inocybe petiginosa o2 5 (o.4,3) .Plo t10 : Cordyceps militaris r25 ,2 (0.1 ,3 ,0.4) , Psilocybe inquilina r25 ,2 (0.1 ,3 ,0.4) .Plo t11 : Rickenella setipes r25 ,3 (0.1 , 3,0.6) , Tubaria conspersa o25 ,3 (0.6,3 ,0.6) .Plo t12 : Lycoperdon spadiceum r25 ,1 (0.1,3 ,0.1) , Cystoderma granulosum o25 ,2 (0.3,3 , 0.3), Ramaria flaccida o25 ,1 (0.3 ,3 ,0.1) .Plo t13 : Cortinarius delibutus r25 ,1 (0.1 ,3 ,0.1) ,Russula nitida r25 ,3 (0.2 ,3 ,0.4) .Plo t14 : Mycena polyadelpha f25 ,8 (1 ,3 ,1) ,Ripartites helomorpha o25 ,3 (0.3 ,3 , 0.4), Russula cyanoxantha f. peltereaui r25 ,1 (0.1,3 ,0.1) , Incrustoporia semipileata r25 , 3 (0.1,3 ,0.4) .Plo t15 : Coprinus stellatus o25 ,3 (0.6,3 ,0.4) , Melanoleuca melaleuca r25 ,1 (0.2 ,3 ,0.1) , Psathyrella cf obtusata o25 ,8 (0.4 ,3 ,1) , Tyromyces floriformis r25 ,1 (0.1 ,3 , 0.1), Hebeloma vaccinum r25 ,1 (0.1,3 ,0.1) , Scleroderma areolatum r25 ,1 (0.2, 3,0.1) .Plo t16 : Cordyceps spec.1 r 25 ,3 (0.1 ,3 ,0.4) .Plo t17 : Coprinus heterosetulosus f25 ,3 (1 ,3 ,0.4) , Collybia cf kuehneriana r25 , 3 (0.1,3 ,0.4) .Plo t18 : Crucibulum laeve o2 0(0.7 ,2) , Helvella crispa o2 0(0.5 ,2) , Russula heterophylla o2 0(0.3 ,2) , Lepiota cristata r2 0 (0.1, 2).Plo t19 : Clitocybe gibba r2 5(0.1 ,3) . Plot20 : Flammulaster carpophiloides r1 7(0.1 ,2) ,Inonotus polymorphus r1 7(0.1 ,2) , Trametes semisupina r1 7 (0.1,2) , Fomes fomentarius r1 7 (0.1,2) .Plo t21 : Daedaleopsis confragosa r2 5(0.1 ,3) .Plo t23 : Russula aeruginea r25 ,4 54 (0.2, 3, 1), Phyllopoprus rhodoxanthus o25 ,2 (0.6,3 , 0.6).Plo t24 : Galerina heimansii o25 , 3 (0.4,3 , 1), Tyromyces subcaesius r25 ,3 (0.3, 3,1) , Mycena hemispaerica 24-XI-1959,le g J.J.Barkma n 6475,det . R.A.Maa s Geesteranus, Russuia fellea found by J.J. Barkman, 22-X-1958 (pers. comm.), Leucocopriaus brebissonii 9-VIII-1960, leg. J.J.Barkma n 6646,det .J.J . Barkman, Clavariadelphus junceus 15-XI-1958,leg .J.J .Bark man 6019, det. R.A.Maa s Geesteranus, Typhula quisquiliaris 24-IX-1960, leg. J.J.Barkma n 6794, det.R.A .Maa sGeesteranus , Macrotyphula fistulosa 15-XI-1958, leg. J.J.Barkma n 6008, det. R.A.Maa sGeesteranus , Phlebia rufa 24-IX-1960,leg .J.J . Barkman 6673, det.M .Donk , Stereum gausapatum 22-X-958, leg.J.J .Barkma n 5840, det.R.A .Maa s Geesteranus (allcollec tions in Herb.WAG-W) . Plot26 : Mycena oortiana o2 5 (0.4,3) .Plo t27 : Pleurotus dryinus r3 3 (0.3,4) .Plo t28 : Psathyrella spec,r 2 5 (0.1, 3). Remarkst oth etable s7 an d8 . Coprinus section Micacei ismentionne di nbot htables .Thi swa sth especie s C. domesticus inth eplot s2 0an d26 ,C . micaceus inplo t24 ,C . radians in plot29 ,an d C. xanthothrix inth eplot s2 5an d28 .I nplo t2 4C . domesticus was also found (J.J.Barkman , 6687, 24-9-1960,det .C .Bas ,Herb .WAG-W) , without indication on abundance.A s the species areclosel y related and have apreferenc e forth eViolo-Quercetu m ilicetosum, they are joinedt o section Micacei in the fungus tables. The species themselves have the followingvalue s forP ,GAMAC ,MAF,'an dMTS Fi ntabl e8 :C . domesticus in Violo-Quercetum typicum I,o.l , 2,0.4 , inViolo-Quercetu m ilicetosum I, 0.1,3 ,- ;C . micaceus I,7 ,3 ,0.3 ; C. radians I,28 ,2 ,- ; C. xanthothrix II,0. ,6 ,- . Cystoderma longisporum hasbee ninclude dwit h C. amianthinum, becausei t wasno teas yt odistinguis hthes especie smacroscopicall y andthe ywer eno t always studied microscopically. C. longisporum was found for certain in plot3 (r17 ,1 )an di nplo t4 (r25 ,3 )an dth evalue swer efo rP ,GAMAC , MAF, and MTSF, inth eDicrano-Quercetu m I,0.1 , 1, 0.5; inth eQuerco - BetuletumI ,0.4 ,3 ,0.6 . Bjerkandera adusta and Polyporus ciliatus werebot h found once inth e Dicrano-Quercetum by P.Ypelaar , in aplo ttha twa sonl yinvestigate dfo r oneyear .So ,bot hspecie sha da A Fo f100% ,and ,mathematically ,a MA Fo f 9%.Th especie sar eno tfrequen ti nth eDicrano-Quercetum ; aMA Fo fI woul d bemor eappropriate . Phellinus ferreus wasals o foundonc ei na Dicrano-Quercetu mb yP .Ypelaa r Ash edi dno tindicat eabundance ,I ca nonl ygiv eth evalue s forP an dMA F intabl e8 . 55 4.3 RESULTS 4.3.1 Number of species Inth e 4 (sub)associationstogethe r 313 fungusspecie s (includingsub species andvarieties )wer e observed.Th e distribution of the speciesi n the 4vegetatio n types (table9 ) shows thatth e total number offungu s species isremarkabl y lowi nth eViolo-Quercetu milicetosum .Thi si sals o expressed inth emea nnumbe rpe rplot .Thes elo wnumber swer emos tlikel y causedb yth esmal lan dno trepresentativ eplo tarea si nth eViolo-Quercetu m ilicetosum. Themea nnumbe ri nth efrequenc yplot si nthi ssubassociation , whichar elarger ,s omor erepresentative ,an dmor eintensivel yinvestigated , dono tdiffe rfro mtha to fth eViolo-Quercetu m typicum.Th eViolo-Quercetu m typicumha sslightl yles sfungu sspecies ,tota lan dmea npe rplot ,tha nth e Querco-Betuletum. Inth eDicrano-Quercetu mth etota lnumbe ro ffungu sspe cies dono t differmuc h from that inth e Querco-Betuletum andth eViolo - Quercetum typicum.Th emea nnumbe rpe rplo tan dpe rfrequenc yplo ti sre markablyhighe ri nth eDicrano-Quercetu m thani nth eothe rvegetatio ntypes , or inothe rwords ,th eDicrano-Quercetu m plotswer ever y richi n fungus species. Table9 :Numbe ro ffungu sspecie spe r(sub)association . DQ QB VQt VQi meanpe rplo t 106.3 76.5 66.9 54.5 meanpe rfrequenc yplo t 115.5 81.6 74.7 74.6 total 179 187 175 136 foundi nal l4 type s 67 67 67 67 foundi nonl y3 type s 20 20 20 3 3 3 30 30 30 foundi nonl y2 type s 24 24 5 5 3 3 11 11 6 6 7 7 foundi nonl yon etype s 57 57 26 35 20 4.3.2 Differential species 4.3.2.1 Introduction Any quantitative parameter canb e used todetermin eth eecologica lan d 56 sociological optimum of aspecies .Her e thevalue s ofP ,GAMAC ,an dMTS F (table8 )wer euse d forthi spurpose .Specie swit hon eoptimu mo rsevera l optimaar efaithfu l (character)species ,an ddifferentia l speciesrespecti vely.Decidin g if aspecie sha sa noptimu mi non eassociatio n (orallianc e etc.)only , isonl ypossibl ei fal lplan tcommunitie so fa regio nwer ein vestigated.Wit h regard tofung iw ear estil lfa rfro mthi sstat eo fknow ledge in anyregion .Therefor e this studyonl ypermit sth edistinctio no f differentialspecie s fora certai nsub-association ,association ,o rallianc e within the group of acid oakwoods.Thi sdoe sno texclud eth epossibility , however, that some of the differential species may turnou t later tob e characterspecies . Aspecie si sconsidere dpresumabl ydifferentia li fi ti sdifferentia li n onecharacteristic ,an di sconsidere ddifferentia li fi ti sdifferentia li n two characteristics and theothe r characteristic doesno tcontradic tit . Themai ndifferentia lcharacteristi c isth eP value .A specie si sconsidere d differentiali fi tha sa P o fIV ,V ,o rV Ian da P a tleas t4 classe slowe r inever yothe rvegetation ,i fi tha s aP ofVII ,VIII , orI Xan da P a t least 5classe s lower inever y othervegetation ,o rwit ha P o fX an da P atleas t6 classe slowe ri never yothe rvegetation . The second differential characteristic isth eGAMA Cvalue .A specie si s considered differential ifth eGAMA C is atleas t2 classe slowe ri never y othervegetation .Th e classificationo fabundanc evalue s (p.48 )wa sused ; 0.1-0.9wa sadde da sa separat eclass -. The third differential characteristic isth eMTS Fvalue ,whic hha sles s distinguishingvalu etha nth eP an dth eGAMAC ,becaus ei twa sbase do nonl y a fewplot s andwa s notmeasure d inal l species.A specie si sconsidere d differential ifth eMTS F is at least2 classe slowe ri never yothe rasso ciation.Th efollowin gclassificatio nwa sused :0.1-0.9 ,1-9 ,10-25 ,26-60 , 61-100. Thepresenc e ofth edifferentia lspecie si nothe rassociation swa sals o studied in literature, inorde r to getsom einformatio no nth eecologica l range. Literature was studiedo naci doakwood s (Quercionrobori-petraeae) , on richer oak andbeec hwood s (Querco-Carpinetum,Fago-Quercetum ,Luzulo - Fagetum,Melico-Fagetum )an d onpoo raci dconiferou swood so rscrub s(Vac - cinio-Piceetalia). 4.3.2.2 TheDicrano-Quercetu m Many ofth e species ofth eDicrano-Quercetu m (tables7 an d8 ,grou p1 ) aredifferentia lo rpresumabl ydifferentia l species,indicate dwit hDQ ,an d pDQrespectivel y intabl e8 .Fourtee nspecie swit ha lo wpresenc etha twer e notdifferentia l according to the criteria,bu twer eno t foundi non eo f theothe rassociations ,wer eadde dals ot ogrou p1 . There are 30differentia l species: Amanita fulva, Boletus edulis, B. 57 erythropus, Cantharellus cibarius, Cordyceps canadensis, C. ophioglossoides, Cortinarius alboviolaceus, C. bolaris, C. fusisporus, C. glandicolor, C. obtusus, C. paleaceus, C. cf stemmatus, Dermocybe cinnamomeolutea, Entoloma turbidum, Hydnellum scrobiculatum, H. spongiosipes, Inocybe napipes, I. ovatocystys, I. sambucina, Lactarius chrysorheus, Leotia lubrica, Marasmius androsaceus, Psathyrella fulvescens var. dicrani, Russuia adusta, R. fragi- lis, Sarcodon scabrosus, Telephora terrestris, and Tricholoma portentosum. Galerina calyptrata is alsodifferentia l forth eDicrano-Quercetum ,bu ti s listed ingrou p7 becaus ei ti sals odifferentia lfo rth eQuerco-Betuletu m inrelatio nt oth eViolo-Quercetum .Presumabl ydifferentia l areth efollow ing9 species : Hebeioma pumilum, Inocybe xanthomelas, Russula cyanoxantha, R. vesca, Tricholoma columbetta, T. virgatum (group1) ,an d Amanita citrina, Laccaria amethystina, and Lactarius camphoratus (listedi ngrou p7 abecaus e they are also differential or presumably differential for theQuerco - Betuletumi nrelatio nt oth eViolo-Quercetum) . No literaturewa s available on fungio fth eDicrano-Quercetum .Man yo f the 'specieso fth eDicrano-Quercetum 'wer eals oreporte d fromothe rasso ciations,sometime so nriche rsoil ,an dofte nbot hfro mdeciduou san dconi ferouswoods .Fo r instance Russula fragilis wasreporte d froma Querco-Be tuletumtypicu m (Runge1960 ,an di nthi sstudy) , froma Genist otinctoriae - Quercetumpetraea esubcarpaticu mdicranetosu m (Bohus& Babo s 1967), aQuerco - Carpinetum (Eihellinger1964 ,Smard a 1972), aLuzulo-Fagetu mleucobryetosu m (Jahn,Nespia k &Tiixe n1967 ,the yfoun di ta differentia lspecie sfo rthi s sub-association),a Melico-Fagetu m (Lange 1978), andals ofro massociation s withconiferou strees ,a Melampyro-Abietetu mvarian twit hLeucobryu mglaucu m (Krieglsteiner1977) , aQuerco-Piceetu m (Nespiak 1959), anda Pino-Quercetu m serratuletosum (Nespiak 1959). Alldifferentia lspecie sar especie so facid ,sand ysoils ,poo ri nhumus . Many ofthe m havea wide recologica lrang etha nth eDicrano-Quercetum ,bu t iti spossibl e thatwhe n growing inriche rwoods ,the yar erestricte dt o poorer,mor e acidpatches .Th e realoptimu m forsom eo fthes especie si s probably inconiferou s woods or scrubs,o ro npoo rheat h lands,suc ha s Marasmius androsaceus and Inocybe lacera. Within the oakwoods onsand y soils inDrenthe ,however ,the yhav e anoptimu m inth eDicrano-Quercetu m andca nb econsidere da sdifferentia lspecies . A few species are possibly character species, Cantharellus cibarius, Cordyceps canadensis, C. ophioglossoides, Hydnellum scrobiculatum, H. spongiosipes, Inocybe sambucina, Psathyrella fulvescens var. dicrani, Sarcodon scabrosus, S. underwoodii. Cantharellus cibarius and the two Cordyceps species are sometimes found outside theDicrano-Quercetum , but theecologica lrang ei snarro wenoug ht oconside rthe ma scharacte rspecies . Inocybe sambucina and the Hydnellum and Sarcodon speciesar erare ,an di n Drenthe only found in theDicrano-Quercetu m (see alsoYpelaa r inprep.) . Sarcodon underwoodii wasmentione donl yonc e (seeanne xt otable s7 an d8) , 58 but itwa s also found inanothe rDicrano-Quercetu m inDrenth e (Ypelaari n prep.)an dwa sno treporte d fromothe rlocalitie si nthi sprovince . Psathy- rella fulvescens var. dicrani wasfoun dofte ni nth eDicrano-Quercetum ,an d hasno tye tbee nreporte d fromothe rhabitats . Many of the specieso fgrou p1 ar emycorrhiza lspecies ,4 0 (=82%).Onl y 6 species (12%)ar ehumu s saprophytes,2 specie sar eparasitic ,1 specie s growso nfalle nleaves .Ther ear en ospecie si nthi sgrou pgrowin go nwood . 4.3.2.3. TheQuercio nrobori-petraea e The Quercion robori-petraeae hasman ydifferentia l species (indicatedwit h Qrpbefor e thenam *o fth especie si ntabl e8 )an dpresumabl ydifferentia l species (indicated withpQ )(grou p2 i ntable s7 an d 8). Somespecie stha t areno tdifferentia lbu thav e apreferenc efo rth eQuercio nwer eadde dt o this group,the ywer e absent from theDicrano-Quercetu m orha dther emuc h lower P orGAMA Cvalues .Th e species were listed insub-group saccordin g preferencefo rth eQuerco-Betuletu m (sub-group a), theViolo-Quercetu m(b) , the Violo-Quercetum typicum (c),th eViolo-Quercetu m ilicetosum (d),th e Querco-Betuletum andViolo-Quercetu m ilicetosum (e),th e Querco-Betuletum andViolo-Quercetu m typicum (f), andwithou ta preferenc e(g) . There are 29differentia l species: Bjerkandera adusta, Chondrostereum purpureum, Clitocybe fragrans, C. metachroa, Collybia cookei, Cudoniella acicularis, Galerina cinctula, Hapalopilus rutilans, Hohenbuehelia atroca- erulea, Kuehneromyces mutabilis, Marasmiellus ramealis, Mycena epipterygia, M. haematopus, M. sepia, M. stylobates, M. vitilis, Oudemansiella platu- phylla, Panellus serotinus, Phallus impudicus, Pluteus salicinus, Polyporus brumalis, Psathyrella hydrophilla, Psilocybe crobula, Stereum hirsutum, S. rugosum, Tephrocybe tylicolor, Tyromyces chioneus and Xylaria hypoxylon. The following 14specie s arepresumabl y differential: Clitocybe diatreta, Clitopilus hobsonii, Crepidotus pubescens, Galerina hypnorum, G. sahleri, Hypholoma sublateritium, Inonotus radiatus, Laccaria laccata, Merulius tremellosus, Mucena mucor, M. pura, Piptoporus betulinus, Pleurotus ostrea- tus, and Rickenella fibula. The other species of this group arespecie s with alo wpresence ,whic hhav e apreferenc e forth eQuercio nbu tar eno t differential. TheQuercio n iswel lcharacterize di nrelatio nt oth eDicrano-Quercetu m andth enumber so fdifferentia lspecie sindicat eth esmal laffinit ybetwee n theQuercio nan dth eDicrano-Quercetum .Th edifferentia lspecie sare ,gener ally,specie so fdeciduou swood so nnutrien tan dhumu sriche rsoil .Man yo f thespecie swer erecorde di nliteratur e fromassociation ssuc ha sth eQuerco - Carpinetum, the Mercuriali-Fagetum, theMelico-Fagetum , e.g. Bjerkandera adusta, Clitocybe fragrans, Mycena haematopus, Xylaria hypoxylon. Someo f the specieswer e recorded fromconiferou swood stoo ,e.g . Collybia cookei, Mycena epipterygia, Laccaria laccata, andon especies , Clitocybe diatreta, 59 was recorded from coniferous ormixe d woods only andno tfro mth eriche r deciduouswoods . Therewer e6 3specie si ngrou p2 ,o fwhic h4 (=6%)wer emycorrhiza lspe cies,1 1specie s (17%)humu ssaprophytes ,3 2specie s (51%)growin go nwoo d andbranches ,1 5specie s (24%)growin go nfalle nleaves ,fruit so rmosses , and 1specie s growingo ndung .Thi s spectrumi scompletel ydifferen tfro m thato fth especie so fth eDicrano-Quercetu m (group1) . The associations ofth eQuercio nhav e distinclyles sdifferentia lspe cies.Th eQuerco-Betuletu m (group3 an dgrou p7 partly )ha sonl y2 differ ential species (indicated withQ B intabl e8) , Mycena rorida and Hymeno- scyphus epiphyllus, and 2presumabl y differential species (indicatedwit h pQB), Panellus stipticus and Clitocybe brumalis. Somespecie stha toccurre d inbot hth eDicrano-Quercetu m andth eQuerco-Betuletu m (thereforeliste di n group7 )ar e alsodifferentia lfo rth eQuerco-Betuletum , Laccaria amethys- tina, Galerina calyptrata, and Lactarius camphoratus, orpresumabl ydiffer ential, Amanita citrina. Cortinarius orellanoides, Galerina heterocystys, and G. triscopa areno t differential according toth e criteria,an dwer e addedt ogrou p3 becaus etheir eabsenc efro mth eothe rassociations . Runge (1960)an dBirke n (1976)gav ebot ha lis to fspecie sfoun di nth e Querco-Betuletum.Rung ementione do fthes especie sonl y Laccaria amethysti- na, Birken mentioned only L. amethystina and Panellus stipticus, soth e similarity withm y study islow .O fth e (presumably)differentia lspecies ' ofgrou p3 Mycena rorida wasrecorde di nliteratur e fromth eQuerco-Carpine - tum (Runge1963) , from 'acidophilousoak-beec kwoods ' (Lisiewska 1974),an d also from coniferouswood s orscrub s ormixe dwoods ,th ePino-Vaccinietu m myrtilli,Pino-Vaccinietu m uliginosi, and Querco-Piceetum (Nespiak 1959), the Melampyro-Abietetum, and the Asperula-Abieti-Fagetum (Krieglsteiner 1977)an dth eDicrano-Juniperetu m (Barkman,pers .comm.) . Clitocybe brumalis is probably more a species ofconiferou swoods ,Krieglsteine r foundi t characteristic for theMelampyro-Abietetu m varianto f Leucobryum glaucum, Nespiak recorded it from thePino-Quercetu m serratuletosum and thePino - Vaccinitummyrtilli ,i ti scommo ni nth eDicrano-Juniperetu m (Barkman,pers . comm.), and itwa sno trecorde db yth eauthor sstudie d fromothe rtype so f deciduouswoods .I di dno tfin drecording so f Galerina calyptrata and Hyme- noscyphus epiphyllus, probably they are lacking there,bu ti ti spossibl e thatthe yhav ebee n overlooked by their small size ormistake nfo rothe r species.O fth e7 specie so fgrou p3 ,ther ei son emycorrhiza lspecies ,on e humus saprophyte, and 5specie sgrowin g ondecayin gwood , leaves,twig s etc. TheViolo-Quercetu mha s3 differentia l species, Collybia fusipes, Mycena inclinata, and Tyromyces caesius, and 2presumabl y differentialspecies , Galerina ampullaceocystis, and Stropharia aeruginosa (group4 ,indicate d 60 withVQ , pVQ respectively). Mycena inclinata has a slightpreferenc efo r thesub-associatio nilicetosum ,th eothe r3 specie s forth esub-associatio n typicum.Th evalue s forP i n Collybia fusipes ishighe ri nth esub-associa tion typicum than inth e sub-association ilicetosum, but thevalue s for GAMAC andMTS F arenearl y thesame . Entoloma euchrous, Inonotus dryadeus, and Russula parazurea wereadde dt ogrou p4 becaus ethe ywer eno tfoun di n oneo fth eothe rassociations . Idi dno tfin drecording so f Galerina ampullaceocystis inth eliteratur e studied.Th eothe r (presumably)differentia lspecie swer eal lrecorde dfro m the Querco-Carpinetum (Nespiak 1959,Einhellinge r 1974, Lisiewska 1965), and also from associations on richer soil, such asth eMelico-Fagetu m (Lisiewska 1974,Lang e 1978)an d the Fagetum typicum (Smarda 1972).The y wereno trecorde d from themor e humuspoo rassociation ,Querco-Betuletum , and can- excep t for Tyromyces caesius -b eregarde da sspecie so fdecidu ouswood s onno tto onutrien tan dhumu spoo rsoils ,bu ti ti sno tknow ni n which association theymigh thav e the optimum. T. caesius certainlyreac h itsoptimu m inconiferou swoods .I tusuall ygrow so nwoo do fconifers ,an d wasrecorde dfro mPinus-Robini awood s (Winterhoff1976) , fromth eMelampyro - Abietetum and Asperulo-Abieti-Fagetum (Krieglsteiner 1977), from theDi - crano-Juniperetum (Barkman,pers .comm.), andals ofro mth eQuerco-Carpine tuman dth eMelico-Fagetum .Coniferou stree swer eno tpresen ti nth eViolo - Quercetum plots of thisstudy ,bu tsometime slog so fconifer swer epresen t (artificial). T. .caesius growing onwoo d of deciduoustrees ,however ,wa s alsoobserved . Of the 8specie s ofgrou p4 ther ei son emycorrhiza lspecies ,on ehumu s saprophyte,an d6 specie sgrowin go nwoo do rbranches . The sub-associations arebot hwel lcharacterize db ya numbe ro fdiffer ential species. The sub-association typicum has 3differentia lspecies , Collybia peronata, Marasmius splachnoides, M. epiphylloides, and2 presum ably differential species, Clitocybe flaccida and Scleroderma verrucosum (group5 , indicated withVt ,an dpV trespectively) . Laetiporus sulphureus, Mycena pearsoniana, Resupinatis applicatus, and Xerocomus parasiticus were added to this group,becaus e theywer e only found inth eViolo-Quercetu m typicum. Collybia peronata was also frequent inth eothe rsub-associations ,bu t herereache dth ehighes tvalue sfo rP ,GAMAC ,an dMTSF .I twa sals orecorde d from other associations, such as theQuerco-Carpinetu m (Lisiewska1965 , Runge1963 ,Einhellinge r 1964),Mercuriali-Fagetu m (Lisiewska 1974),Melico - Fagetum (Lisiewska 1974,Lang e 1978), Melampyro-Abietetum (Krieglsteiner 1977), and the"Dicrano-Juniperetu m (Barkmanpers .comm.) . Jahn,Nespia k& Tüxen(1967 )calle di ta accompanin g speciesi nth e Fayus woodsthe yinves tigated. Marasmius splachnoides hasabou tth esam evalu efo rGAMA Cher ea s inth eQuerco-Betuletum ,bu tmuc hhighe rvalue sfo rP an dMTSF .I twa sals o 61 recordedfro mth eQuerco-Carpinetu m (Lisiewska1965) ,th eMercuriali-Fagetum , and the Melico-Fagetum (Lisiewska 1974). Also from ownexperienc e iti s known to occur inriche r oakwoods sucha sth eQuerco-Carpinetum ,wher ei t possiblyreache sit soptimum . Marasmius epiphylloides isconfine dt oterres trialgrowin g Hedera helix, andtherefor erestricte dt oth eViolo-Quercetum . Itca nprobabl yb ea characte rspecie si nth eNetherlands .I nliteratur ei t was recorded only from theQuerco-Carpinetu m (Lisiewska 1965). Idi dno t find data inliteratur eo nth edistributio no f Clitocybe flaccida. There lated taxa C. gilva and C. inversa wererecorde d fromth eQuerco-Betuletu m (Birken 1976), theQuerco-Carpinetu m (Runge 1963,Einhellinge r 1964)an d theLuzulo-Fagetu m leucobryetosum (Jahn,Nespia k& Tiixe n1967) . Scleroderma verrucosum isrecorde d fromth eQuerco-Carpinetu m (Lisiewska 1965), andth e Melico-Fagetum (Lange 1978). Thespecie so fthi sgrou par eal lspecie soc curring inriche r oakwoods ,ofte nals oi nbeec hwoods .Thei rabsenc efro m theViolo-Quercetu m ilicetosumi sprobabl ycause db yth eunfavourabl elitter , owingt oth elarg eproportio no fIle xleaves ,o rb yth eunfavourabl emicro climate. Of the 9specie s of thisgrou p there ison emycorrhiza l species,on e parasitical species,an d7 specie sgrowin go nhumus ,litte ro rwood . Thesub-associatio nilicetosu mha s6 differentia ltaxa , Ciboria batschi- ana, Coprinus section Kicacei, Polyporus varius, Psathyrella frustulenta, Typhula erythropus, and T. phacorrhiza, and2 presumabl ydifferentia lspe cies, Cortinarius tabularis and Mutinus caninus (group6 , indicatedwit h Vi, andpV irespectively) . Phellinus ferreus wasals oplace di nthi sgroup , because itwa s found only once in one of theothe rassociation s (without data on abundance or exact habitat). Four species of Coprinus section Micacei were found,C . domesticus, C. micaceus, C. radians, andC . xantho- thrix, which allha d apreferenc e forth eViolo-Quercetu m ilicetosum,bu t the individual presence degreeswer elow .Th esectio na sa whole ,however , isdifferential .Th e speciesals ooccu ri nothe rassociations .C . micaceus was recorded from theQuerco-Betuletu m (Runge 1960),th eQuerco-Carpinetu m (severalauthors) ,a Pinus-Robinia wood (Winterhoff1977) ,th eMelico-Fage tum (Lange1978 ,an dJahn ,Nespia k& Tiixe n1967 ,wh ocalle di ta presumabl y differential species forth eMelico-Fagetum) .C . xanthothrix wasrecorde d from the Querco-Carpinetum (several authors), and from Pinus-Robinia and Teucrium-Quercus-Pinus woods (Winterhoff1977) , C. domesticus froma Pinus- Robinia wood (Winterhoff 1977). Invie w ofthei rhabitat ,rotte nwoo dan d twigs ofdeciduou s treesI expec ttha tthe yhav erathe rwid erange sa sfa r as wood associations are concerned. Both Typhula specieswer e foundt o occur only inthic klayer so f Ilex leaves (seep .87) .The yar eals oknow n fromlitte ro fothe rdeciduou strees ,bu tusuall yi na fairl ymois thabitat . Polyporus varius wasrecorde d fromth eMelico-Fagetu m (Lange 1978), Mutinus caninus from the Melico-Fagetum (Lange 1978) and the Querco-Carpinetum 62 (Lisiewska1965) . Idi dno tfin ddat ai nth eliteratur eo nth edistributio n inothe r associations of the other (presumably)differentia l species.Th e number ofdifferentia l species of themycocoenos eo f theViolo-Quercetu m ilicetosum is large compared with thato fth ephytocoenos e (onlyon edif ferential species). Iti stherefor epossible ,base do nth emycocoenose ,t o consider theViolo-Quercetu m ilicetosum as anindependen tassociatio nan d nota sa sub-association . Of the 12specie s ofgrou p6 ,ther ei son emycorrhiza lspecies ,2 humu s saprophytes,3 specie s growing on fallen leaves andfruits ,an d6 specie s growingo nwoo do rbranches . Therear e1 9specie swit ha preferenc e forth eDicrano-Quercetu m andth e Querco-Betuletum (group 7). Listedwer especie stha twer edifferentia lfo r both associations in relation to the Violo-Quercetum, and species that occurredwit ha lo wpresenc ei nth eDicrano-Quercetu m andth eQuerco-Betule tuman dwer eabsen ti nth eViolo-Quercetum . Sevenspecie shav ea preferenc e forth eDicrano-Quercetu m (sub-group a), 4o fthe mar e (presumably)differ ential forth eDicrano-Quercetu m orth eQuerco-Betuletu m (seethere) .Twelv e species occurred almost equali nth e2 associations ,wit hlo wpresenc ean d abundance, andwer e listed as 'indifferentspecies ' (sub-group b). These species were all absent inth eViolo-Quercetum .Ther ewer en ospecie sbe longing to this group thatha d apreferenc e forth eQuerco-Betuletum .Ap -. parentlythes especie sca nb econsidere d asspecie so f (oakwood son )acid , sandy soils,poo r inhumus .E.g .bot h Laccaria species whenoccurrin gi n theViolo-Quercetum ,prefere dsite swit ha sand ylaye ra tshallo wdepth . There were 23specie s present in the Dicrano-Quercetum, theQuerco - Betuletum,an dth eViolo-Quercetu m typicum,an dabsen to rpresen twit hmuc h lowerP andGAMA Cvalue si nth eViolo-Quercetu m ilicetosum (group 8). They wereliste di nsub-group saccordin gth epreferenc e forth eDicrano-Quercetu m (sub-group a), forth eDicrano-Quercetu m andQuerco-Betuletu m (b), forth e Querco-Betuletum andth eViolo-Quercetu m typicum (c), forth eDicrano-Quer cetum and theViolo-Quercetu m typicum (d), andwithou tpreferenc e (e).Th e specieswit ha preferenc e forth eDicrano-Quercetum ,o rth eDicrano-Querce tum and theQuerco-Betuletum , resembled the specieso fgrou p7 ,bu tha da wider range, Russvla emetica, Cystoderma amianthinum, Collybia cirrhata, Xerocomus badius, and Galerina atkinsoniana. Collybia butyracea and Clito- cybe candicans had apreferenc e forth eQuerco-Betuletu m and theViolo - Quercetum typicum.Ther ewer e5 specie sliste di nsub-grou pd ,bu tth epref erence forth eDicrano-Quercetu m and theViolo-Quercetu mwa sonl yweakly , because thevalue s forP ,GAMAC ,an dMTS Fwer ever ylow .Th eothe r1 1spe cies ingrou p8 wer emor e or lessequall ydistribute di nth e3 vegetatio n types. The absence ofthes especie s fromth eViolo-Quercetu m ilicetosumi s acharacteristi c theyhav ei ncommo nwit hth especie so fgrou p2f .Probabl y the forest floor in theViolo-Quercetu m ilicetosum was unfavourable for 63 these fungi becauseo fth e largeproportio no fdry ,brittle ,an dbadl y decomposing Ilex leaves andb yth eabsenc eo fmycorrhiz ai nIlex. Thedif ferencesi nmicroclimate ,cause db yth edens eshad ethroughou tth eyear , or byth eabsenc eo fth eher blayer ,migh tb eanothe rfactor . Accompanying species (group9 ,3 8 species) areth especie s thatwer e moreo rles s equally distributed troughoutth etable .Liste dher ear eth e very common species,wit hhig hvalue sfo rP ,GAMAC ,an dMTSF ,bu tals oth e rarer specieswit h lowvalue s forP ,GAMAC ,an dMTSF .Som eo fth especie s were most abundanti nth eDicrano-Quercetum , e.g. Lactarius quietus, I. theiogalus, Paxillus involutus, inth eQuerco-Betuletum ,e.g . Mycena san- guinolenta, ori nth eViolo-Quercetu m typicum,e.g . Clitocybe vibecina, and Armillariella mellea. Thecommo n species sometimesha dGAMAC .MTSF ,o rP values thatwer e differential accordingt oth ecriteria .Bu tbecaus eth e valueswer e highi never yvegetatio n type,thes e specieswer eno treall y differential,an dtherefor eliste di nthi sgroup .Generally ,th especie os f thisgrou par especie swit ha wid erange ,occurrin gals ofrequen ti nothe r associations. 4.3.3 Mathematical affinities Thevalue s forA c,Ap ,an dA- ,(tabl e11 )an dfo rD , D p,an dD _ (fig.9 , the lengtho fth econnection si sa measur e fordifference s betweenth e associations)sho wtha tth eaffinitie so fth efungu svegetatio nar emor eo r lessth esam ea sthos eo fth evegetatio no fphanerogams ,mosse san dlichens . Therei sa grea t affinity betweenth eViolo-Quercetu m typicuman dth e Querco-Betuletum,eve ngreate rtha nbetwee nbot hth esubassociation so fth e Violo-Quercetum. Theaffinit yo fth e Dicrano-Quercetum withth e other vegetationswa sver y low,especiall y withth eViolo-Quercetu milicetosum . Thesam erelation swer eshow ni nth eaffinitie so fth ephytocoenoses . Therelatio no fth eaffinitie sappeare dt ochang ei fno tonl yth enumbe r of specieswa suse d (fig.9a) ,bu tals oth ePresenc e (fig.9b )an dth e GAMACvalue s (fig.9c) ,jus ta si nth eaffinitie so fth ephytocoenoses . But herebot hth eaffinitie so fth eViolo-Quercetu m ilicetosum andth eDicrano - Quercetumt o theQuerco-Betuletu m andth eViolo-Quercetu m typicumdecreased . Especiallyth eaffinit yo fth eDicrano-Quercetu mwit hth eothe rvegetation s isextremel ylo wi nth evalue sfo rA™ . 4.3.4 Representative plot size The relationbetwee n thenumbe ro fspecie si na plo tan dth e sizeo f that plot (fig.10 )wa sstudie dt odetermin eth e'minimal 'are ao rth e representativeplo tsize .Th enumbe ro fspecie sdistinctl yincrease dwit ha plotare aincreasin gfro m30 0m 2u pt o150 0m 2.Th evariatio ni n thenumbe r ofspecie si nplot so f150 0m 2wa srathe rlarge ,5 0t o10 1species ,wit ah n 64 Table11 :Mathematica l affinities according to 3differen tcalculations . Seetext . Ac Ap "m DQ- Q B 1.61 1.14 0.40 DQ- VQ t 1.08 0.90 0.33 DQ- VQ i 0.89 0.62 0.16 QB- VQ t 2.50 3.15 2.36 QB- VQ i 2.18 2.01 1.25 VQt-VQ i 2.27 2.33 1.33 VQt VQi- VQt Fig.9 .Th e differences between the fungusvegetations , aaccordin gth e calculationo fD ,b accordin gth eD p,an dc accordin gth eD_ . 65 Om o o DO > > • ooo • e*• o E <=> Fig.10 .Th erelatio nbetwee nth eare aan dth enumbe ro fspecies . 66 average of 75species . Inplot s of200 0m 2 the numbero fspecie sdi dno t differmuc h from this; thevariation , 68 to 105 species,wa s nearly the same, and the average of 80specie swa s only slightly higher.O f the2 plots of300 0m 2, oneha d 82specie s anddi dno tdiffe rfro mth eplot so f 1500 and 2000m 2. Theothe rone ,a Dicrano-Quercetu mplot ,surveye ddurin g 6years ,ha dmor especies ,namel y126 . Aplo t seems tohav ea representativ e sizei fi tha sa nare ao f150 0t o 2000m 2, exceptperhap s inth eDicrano-Quercetu mwher ea plo to fthi ssiz e had only 83%o f the species in aplo to f 3000m 2. An areao f100 0ra 2i s certainly not large enought ob erepresentative .Thi si sals oshow nb yth e frequencyrelevé s (sizeabou t100 0m 2)havin ga naverag eo fonl y74 %o fth e speciesfoun di nth ewhol eplo t (size120 0- 300 0 m2). The original idea,tha t 1000m 2 wouldb e arepresentativ e plotsize , appeared to be inaccurate. In the associations ofth e Quercionrobori - petraeae the representative plotsiz eappeare dt ob eabou t150 0m 2, inth e Dicrano-Quercetum atleas t300 0m 2. 4.4 CONCLUSIONS Manymor e fungus speciestha nplan tspecie swer efound .Th eViolo-Quer - cetum ilicetosum appeared to be relatively poor, the Dicrano-Quercetum relativelyric hi nfungi . A number ofth e fungus species aredifferentia lspecies ,jus ta sther e aredifferentia l plant species.Mor edifferentia l fungusspecie stha ndif ferential plant species werepresent ,especiall y inth eassociation swit h only a few plant species,th eDicrano-Quercetu m and the Violo-Quercetum ilicetosum. TheDicrano-Quercetu m and theQuercio nrobori-petraea ehav ei nrelatio n toeac h otherman y differential species.Thi s supports the ideatha tth e Dicrano-Quercetum does not belong to the Quercion robori-petraeae.Th e Quercion associations have in relation to eachothe r less differential species,bu tar eals owel lcharacterized .Base do nth edifferentia lspecie s it is possible to consider ranking theViolo-Quercetu m ilicetosum asa n association. The affinities ofth e fungusvegetation s of the (sub)associationsar e comparablewit hth eaffinitie so fth ephytocoenoses . The representative plot size appeared tob e rather large,ca .150 0m 2 inassociation s ofth eQuercio n robori-petraeae, and at least300 0m 2 in theDicrano-Quercetum . 67 5 MUTUAL FREQUENCY Theresearc htoo k4 years ,s oi twa spossibl et odetermin eth enumbe ro f species found in one,two ,three ,an d fouryears ,an d to distinguish 4 Annual Frequency Classes. Ineac hplo tth enumbe r of species inth eA F Classeswa sdetermine d as aproportio no fth etota lnumbe ro fspecie si n theplot ,an dth emea no fthes evalue spe r (sub)associationwer ecalculate d (fig.11) .Dat a from earlier research and the incomplete date fromth e plots6 , 7, 27, and 29 (these plots werevisite d only in3 years )ar e omitted. The graphs dono tdiffe rgreatl yfro meac hother .A FClas s1 i si neac h associationth elarges t one and tends toincreas e slightly fromDicrano - Quercetumt oQuerco-Betuletu mt oViolo-Quercetu mtypicu mt oViolo-Quercetu m ilicetosum. AFClasse s3 and 4 tend todecreas ei nthi sseries .Thes ear e relatively large inth eDicrano-Quercetu m and inth eQuerco-Betuletum ,o r in otherwords ,th eDicrano-Quercetu m and theQuerco-Betuletu mhav ea re lativelyconstan tmycoflora . Table7 give sth evalue sfo rA Fa sa proportio no fth enumbe ro fresearc h years (inpercentage) . Earlierresearc hb yYpelaa ran dBarkma ni sinclude d here.Tabl e8 give spe rassociatio nth eMea nA Fvalues . 12 3 4 1 2 3 4 12 3 4 12 3 4 Dicrano-Quercetum Querco-Betuletum Violo-Quercetum Violo-Quercetum typicum ilicetosum Fig.11 .Annua lfrequencie spe r (sub)associationi nproportio n(mea n percentage)o fth enumbe ro fspecies . 68 6 SPATIAL FREQUENCY 6.1 INTRODUCTION Therea labundanc eo fa specie si si nfac tth eabundanc eo fth emycelia , and not the abundance of thecarpophores .Th e abundance of themycelia , however, isdifficul t to determine.Darimon t (1973)determine s theabun danceo fth e 'stations'.A 'station'i sa grou po fcarpophore stha ti ssep arated fromothe rgroups ,an dtherefor emark sth espo twher eth eindividual , themycelium , grows.Th e following difficulties mayoccu ri nworkin gwit h 'stations'.Th e 'stations'ar eno tsharpl yseparated ;no tal lmyceli afor m carpophores atth e same time;a 'station'ca nb emad eu po f2 mycelia ,o r 2 'stations' are in fact one mycelium. In this studybecaus e ofthes e difficulties theTota l Spatial Frequency was introduced as an indication for the abundance ofth emycelia ;i ti sexpecte dt ob emor eexac ttha nth e abundanceo f 'stations'. The Spatial Frequency of a species depends on the sizeo fth epartia l relevés, ando n the size,th enumber ,an dth espatia ldistributio no fth e individual.Th e size ofmyceli ao fman ymacrofung i (parasitesan dfung io n dung are excluded here)wil lprobabl yb e1 t o2 0m 2 (basedo nth esiz eo f groups of carpophores). Toge tth emos taccurat evalue sfo rS Fth epartia l relevésmus tb e not smaller than themycelia .Onl yi fal lpartia lrelevé s have the same sizeca nw ecompar eth evalue s forth eSpatia lFrequency .I n this study the sizewa s 25m 2. This isslightl y largertha nth eexpecte d sizeo fmos tmycelia .I fa myceliu mwa slarge rtha n2 5m 2 thiswa srecorde d whenth edat awer eworke dout . Themanne r ofgrowin g and therefore thespatia ldistributio no fa spe cies greatly influences theSF .Becaus eo fthi si ti sno tpossibl et ocon vert abundance ofcarpophore s into Spatial Frequency.Becaus eo fth esiz e chosen forth eparia iplots ,i t is expected thatth e SF is amor eexac t method ofestimatin g the abundance ofmyceli a than itwoul d beb y esti matingth eabundanc eo fcarpophores . 6.2 METHODS In order to make frequency relevés 17plot s (DQ2 plots ,Q B5 plots , VQt7 plots ,an dVQ i3 plots )wer edivide di n35 ,40 ,o r4 2partia lplots , depending onth e sizeo fth ecomposit eplot .A clea rpictur eo fth eS Fca n be obtained from thesepartia lplot s and there areno t tooman y fori t becomeunworkable .Onc eo rtwic ei n1976 ,1977 ,an d197 8a frequenc yrelev é 69 wasmad ei neac hcomposit eplot .A frequenc yrelev éconsist so f35 ,40 , or 42partia l relevés.Al lspecie s werenote di neac hpartia lplo tan dth e abundanceo fcarpophore swa sestimated .Th efrequenc yrelevé swer eunite d perplo tint oon e 'totalfrequenc yrelevé' ,whic hindicate di nwhic ho fth e partialplot sth especie sha dbee n found.Th eTota l SpatialFrequenc yi s theproportio no fth eplo t area (inpercentage )i nwhic hth especie swa s observed duringth e3year sresearch .I nthi spape rTota lSpatia lFequenc y willb erefere dt oa sfrequenc yo ra s TSF. Per associationth emea no fth evalue sfo rTSF ,th eMea nTota lSpatia l Frequency (MTSF),wer ecalculated . Ineac hcomposit eplo tth eportio no fspecie swa scounte d (inpercentag e ofth etota l numbero fspecie si ntha tplot )occurrin gi n1-20 %(Frequenc y Class I), in21-40 %( FII) ,i n41-60 %( FIII) ,i n61-80 %( FIV) ,an di n 81-100%( FV )o fth eplo t area,an dth emea nvalue spe r(sub)associatio n werecalculate d (fig. 12). Indiceso fheterogeneit ywer edetermine daccordin g H=(FI+ FII)/( FIV+ FV) . 6.3 RESULTS 6.3.1 Introduction Asth efrequenc yrelevé swer emainl ymad ei nSeptember ,October ,an dth e firsthal fo fNovember ,an donl ya fe wtime si nJune ,Jul yan dAugust , the valuesfo rth eTS Fca nb eto olo wi nspecie swit hoptimum sbefor eSeptembe r or after themiddl eo fNovember .Thi si sonl yapplicabl et oa fe wspecies , sucha s Oudemansiella platyphylla, Mycena cinerella, Clitocybe vibecina. Some other,rare r species were foundi na composit eplo tbu tno tdurin ga frequency relevé,s othei r spatial frequency wasno t measured.Th e TSF- valuear elikel yt ob emor e accuratei nspecie swit ha hig hAF-value . Ingenera lth eTSF-value sshoul db econsidere da sminimu mvalues . Valuesfo rTS Far egive ni ntabl e7 pe rplot ,an dthos efo rMTS Fpe r associationi ntabl e8 . 6.3.2 Frequencyhistograms .and homogneity The frequency histograms foreac ho fth e4 association san dfo rth e 4 associations together (fig.12 )d ono tdiffe ra grea t deal.Frequenc y ClassI i sver ylarge ,muc hlarge rtha nth eothe rClasses .Clas sV i sabou t twicea slarg ea sClas sI Vi nth eassociation sQuerco-Betuletum ,Violo-Quer - cetumtypicu man dViolo-Quercetu milicetosum .Thi sconfirm swha tRaunkiae r (1934)found ,namel ytha tth eClasse sar ei naproportio no fI>II>III B I 1B iE a f o o aï i> — E a 3 O u > - H E TaB 1l6 J_B1 —» O fl-ï laB s.2 Fig. 12. Frequency histograms for each of the four (sub)associations and for the combined associations, in mean percentage of the number of species. 71 Violo-Quercetumilicetosum . A frequencyhistogra mwit h ClassV largertha nClas sI Voccur sonl yi f thevegetatio n ishomogeneou s and ifth epartia lrelevé sar eo fa certai n size in relation toth e 'minimalarea' .Homogeneit yca nb emeasure db yth e indices ofhomogeneit y orb y thereciprocal ,th e index ofheterogeneity . Heterogeneity forth eDicrano-Quercetu m is14.3 ,fo rth eQuerco-Betuletu m 9.3, forth eViolo-Quercetu m typicum8.5 ,an dfo rth eViolo-Quercetu m ili cetosum 9.3. The Dicrano-Quercetum isnotabl ebecaus e ofth e relatively largeheterogeneity . Raunkiaerconsider stha tth esiz eo fth epartia lrelevé si nwood sha st o beca .0.125 %o fth e 'minimalarea ' (minimalare ause dher ei nth esens eo f theusua l relevé sizei nth eBraun-Blanque tmethod) .Th eFrequenc yClas sI getsver ylarg ei fth epartia lrelevé sar eto osmall .Al lfrequencie sgrou p together inFrequenc y ClassV i fth epartia lrelevé sar eo fabou tth esam e sizea sth e 'minimalarea' .Th epartia lrelevé sar e2 5m 2,tha ti s0.83 %o f theexpecte d minimal area in theDicrano-Quercetu m (atleas t3,00 0m 2 for fungi)an d 1.67%i nth eassociation so fth eQuercio n (1,500m 2 forfungi) . FrequencyClas sI i srathe rlarge ,althoug hth epartia lrelevé sar eno tto o small.Frequenc yClas sI appear st odecreas ean dFrequenc yClasse sI Van dV appear to increase ifw e jointhes epartia l relevés together topartia l relevéso f5 0m z (fig.14) . ClassV in the Dicrano-Quercetum, however, is smaller thanClas sIV . Apparently,Raunkiaer sprincipl edoe sno tappl yt oth efung io fth eDicrano - Quercetum.Possibl y thisi sa specia l characteristico fth eDicrano-Quer cetum or itma yb e caused by one ofth efollowing :th evegetatio no fth e plotswa sno thomogeneous ;th epartia lplot swer eto osmal li nrelatio nt o the 'minima area', inothe rwords ,th e 'minimalarea 'o fth eDicrano-Quer cetum ismuc h larger than ofth eothe rassociations ;th erelevé swer ein complete,no t enough relevés and/or the relevéswer emad ea ta nincorrec t time. Frequencyhistogram so fth ehighe rplant san do fth emosse s (fig.13 )d o not indicate heterogeneity for theDicrano-Quercetum . Thedistributio no f mosses inside theplot s shows someheterogeneit y (seep .82) .Thi si sin herent in the Dicrano-Quercetum. The partial relevés are 0.83%o fth e 'minimal' area'i ftha t is3,00 0m 2 large,o r 0.25%i fi ti s10,00 0m 2. When thepartia l relevés are joined together topartia l relevéso f5 0m 2 or 100m 2 the Frequency Classes IV andV increase somewhat (fig.14 )an d the indices ofheterogeneit y decrease from 14.3 to 5.9 to3.1 .However , Raunkiaers principle still does not fit.Perhap sth e 'minimalarea 'i ss o large that thepartia l relevésmus tb e the size ofth emos spatche s (see p.82) .Th enumbe r of frequency relevés and thenumbe r offung ii nthes e frequency relevés is forth eDicrano-Quercetu m neither extremely lowo r highwhe n compared withth eothe rassociations .Th eAnnua lFrequenc yshow s a relatively constantmyco-vegetation , so iti sunlikel ytha tth erelevé s 72 i n m n ï i n m E Ï Fig.13 .Frequenc y histograms for the higher plants (left)an d forth e mosses (right)i nth eDicrano-Quercetum . inmizz ! 1 I I I Fig. 14. Frequency histograms for partial relevés of 50 m2 (left) and of 100 m2 (right) in the Dicrano-Quercetum. 73 areincomplete . The conclusion is thatth eDicrano-Quercetu mplot shav ea heterogeneou s vegetation ofmosse s and fungi.Th e 'minimal area' for fungima yb ever y large.I tshoul db enote dtha tth emyco-vegetatio no fth eDicrano-Quercetu m isheterogeneou s (inspace )an dconstan t (intime) . 6.3.3 The relation between the frequency and the abundance The relationbetwee n theTS F and theabundanc eo fcarpophore s (highest abundance of carpophores measured in the frequency relevés wasused )wa s similar ineac hvegetatio ntyp efo r (fig.15 ,4 plot swer ecompared) .Mos t ofth e fungi inthi s figure,62% ,ha d acharacteristi c abundance of 1-5 specimenspe rpartia lplot ,o fthes e85 %ha da TS Fo f50 %o rless ;13 %ha d a characteristic abundance of 6-10 specimens perpartia lplot ,16 %o f1 1 andmor e specimens,an d 9%o f lesstha n 1specime npe rpartia lplot .Th e lastmentione d were species that always grewi nlo wabundanc ean dSF ,an d reached ahighe r TSFbecaus e they grew indifferen tpartia lplot sever y year. Species with an abundance ofcarpophore s ina composit eplo to f1 0o r lessha d aTS F of 1-22(-32)% ,thos ewit h abundance of11-5 0ha da TS Fo f 1-68(-88)%, and thosewit h 100an d moreha d aTS Fo f15-100% .Becaus eo f this converting the abundance ofcarpophore s intoSpatia l Frequency,o r Spatial Frequency into abundancei simpossible ,o ronl ya ver yroug hesti mateca nb emade . Forspecie stha tgre wi nnearl yth esam eabundanc ei na partia lplot ,i t ismor eo rles spossibl et odetermin ea characteristi c relationTS F- abun dance. Four examples aregive n in fig.16 . Pluteus cervinus hada charac teristicabundanc eo f1-1. 3 specimenspe rpartia lplot ,an da TS Fo f25 %o r less. Phallus impudicus hada characteristi cabundanc eo f1-1. 6 ifth eTS F was low, <15%, and up tonearl y 3 ifth eTS Fwa shigher . In Hypholoma fasciculare this relation is less stable.Th evalue s grouped themselves around 10specimen s per partialplot ,bu t also 60specimen s perpartia l plotwa s reached. In Armillariella mellea theabundanc ewa sfro m1 t omor e than 25specimen s perpartia lplot ,showin gtha ta characteristi crelatio n betweenth eTS Fan dth eabundanc edi dno texist . 74 o a a e E 9 'ü 2m 5 o a IK ui 2o>? o a o o oo a a aa . a a\ • V \••a wo 15 S°\ am -Qq» *- . o m m Jo' a "S •; o 33i Fig.15 .Th erelatio nbetwee n theTota lSpatia lFrequenc yan dth eAbsolut e MaximumAbundanc eo fCarpophore s infou rplots . 75 o _ ö. o ö £• € .2 £ 1 e f mai i i S .S* f « 'S 'M*5 a ft S .i | 5 (5 « £ o S = ° = = lf E a o. i < .§_ Va ^ '"—8 ° £ t/t o <<£>.o Fig. 16. The relation between the Total Spatial Frequency and the Absolute Maximum Abundance of Carpophores in four species. 76 7 DISTRIBUTION PATTERNS 7.1 INTRODUCTIONAN DMETHOD S The total frequency relevéswer e 'mapped' foreas y recognition ofth e spatialpatterns .A 'map'wa sdraw no feac hfungu sspecie si neac hcomposit e plot.Th eAbsolut e Maximum Abundance ofCarpophore swa sindicate d ineac h partialplot :1 ,2 ,an d3 specimen swer ecounted ,4 an dmor especimen swer e divided intoclasse s of4-1 0 specimens, 11-29, 30-79,80-189 ,an d19 0an d morespecimens .Thi si sVa nde rMaarei' s (1971)classification . The distribution patterns of the fungus species in acomposit eplot , studied on the 'maps' could be random, underdispersed or overdispersed (=clustered).Pielo u (1969:107 )give s astatistica lmetho d fordiscrimi nating random fromunderdisperse d and overdispersed ina gridma p foreac h spatialfrequency .O nth emap sth enumbe ro fjoin swer ecounte dbetwee nth e partialrelevé swher ea specie swa sfound .Th edistributio ni scalle dunder - dispersed ifther e are less joins (significant withp<0.05 )tha nwa sex pected according the nullhypothesi so frando mminglin go fpartia lrelevé s withan dwithou tth especies .Th edistributio ni scalle dclustere di fther e aremor ejoin s (significantwit hp<0.05 )tha nwa sexpected . Themap swer e alsouse d to recognise correlated distribution patterns of fungi inrelatio n toeac h other,o r of fungii nrelatio nt omosse so r higher plants. Correlationbetwee nspecie si sexpresse db yth ecoefficien t V=(ad-bc)/(mnrs)% (ai s thenumbe r ofpartia lrelevé swher ebot hspecie s werepresent ,b andc th enumbe ro fpartia lrelevé swher eonl yon especie s waspresent ,d th enumbe ro fpartia lrelevé swher ebot hspecie swer eabsent , m=a+b,n=c+d , r=a+c,r=b+d. )V can reachvalue sbetwee n- 1an d+1 .I twa s tested according to Pielou (1969: 163)whethe r the values for Vwer e significant.I fsignifican twit hp<0.0 5the na * i splace dafte rth evalue s forV .Value s forV aremeaningles s forspecie swit ha ver ylo wTSF ,<5% , ora ver yhig hTSF ,>90% . Correlation canb e expectedi nfung itha tdepen do nothe rfungi ,mosse s orhighe rplants .I ngenera la plo ti sheterogeneou s fora dependen tfungus , becauseth esubstrat ei susuall yno thomogeneousl ydistributed . 7.2 DISTRIBUTIONPATTERN S Threepattern sar epossible ,rando m (actually:no tsignificantl ydiffer entfro mrandom) ,significantl y clustered,an dsignificantl yunderdispersed . Significantly underdispersed patterns were not found atall .Apparentl y 77 underdispersion isno tonl y scarse inhighe rplant sbu ti ti sals oscars e infungi .Significan tclusterin gdi dno tofte noccur .68 0distributio npat terns were observed; thepatter nwa srando m 537time s and clustered 143 times.Mos tspecie s (109)wer erando mi nal lo ri nal lbu ton eo fth eplot s inwhic hthe ywer efound .Onl y3 5specie swer eclustere di n2 o rmor eplots , of which only 10specie s were clustered rather thanrando m distributed: Amanita rubescens (random 1*,clustere d 2*), Clitocybe flaccida (r0 ,c 2 ) (fig.17) ,Galerina calyptrata (r1 ,c 2), Hapalopilvts rutilans (r1 ,c 2) , Inocybe napipes (r1 , c 2), Lactarius chrysorheus (r0 ,c 2), Marasmius epiphylloides (r0 ,c 3), Scleroderma citrinum (r4 ,c 7 )(fig .18) , Typhula phacorrhiza (r0 ,c 2), and Tyromyces chioneus (r1 ,c 3). Sixspecie swer e as often clustered as random: Collybia peronata (r 3, c 3) (fig. 19), Gymnopilus hybridus (r3 ,c 3), Laccaria amethystina (r2 ,c 2), Lactarius camphoratus (r2 ,c 2), Polyporus bmmalis (r2 ,c 2), and Psilocybe crobula (r2 , c 2). Thw other 19specie s weremor e often random thanclustered : Armillariella mellea (r7 , c 2), Calocera cornea (r12 ,c 3), Collybia butyracea (r10 ,c 3), Collybia cirrhata (r4 ,c 3), Cudoniella acicularis (r10 ,c 2), Hypholoma fasciculare (r14 , c 3), Hypholoma sublateritium (r4 ,c 2), Lactarius quietus (r9 ,c 5), Lactarius theiogalus (r10 ,c 3) , Marasmius androsaceus (r3 ,c 2), Mycena cinerella (r6 ,c 2), Mycena rorida (r5 ,c 2), Mycena sanguinolenta (r10 ,c 2), Mycena stylobates (r5 ,c 2) , Nectria sp. (r11 , c 3), Oudemansiella platyphylla (r9 , c 4), Paxillus involutus (r10 , c 3), Phallus impudicus (r7 , c 3), and Psathyrella squamosa (r10 ,c 4) . Generally the fungus species were onlyclustere d in a fewplots .Onl y Scleroderma cirtinum, Lactarius quietus, and Psathyrella squamosa formed patches in 4o r more composite plots.Patche s canb e formed ifa .th e mycelium is soextende dtha ti ti ssprea dove r2 o rmor epartia lplots ,o r b.th emyceli aar eclustered . a.Th emyceliu m was spread over severalpartia lrelevés .I tma yhappe n thatcarpophore stha tapparentl ybelon gt oa singl emyceliu mgrowin ga tth e bordero fa partia lplot ,ar esprea dove r2 o rmor epartia lplots .Thi swa s often observed in fungi growing on prostrate decaying stems, such as Hapalopilus rutilans, Tyromyces chioneus, Polyporus brumalis, and Gymnopilus hybridus. (fig.20-23) . A smallpatc hwil lprobabl yb eforme db yonl yon e mycelium. Itwa s notpossibl e to determinei nextende dpatche swhethe ri t was one extended mycelium,o rsevera lsmal lmyceli agrowin gtogether .E.g . thecluster si n Clitocybe flaccida inplo t1 2 (fig.17) , Scleroderma citri num inplo t1 4an d 20 (fig.18) , Collybia peronata inplo t1 6 (fig.19) . Thismake si tdifficul tt oseparat epoin ta fro mpoin tb . b.Th emyceli aar eclustered . Ifth eclusterin go fth emyceli ai scause d by intrinsic growth characteristics of the species,I expecte d them to cluster in all theplot s inwhic hthe ywer eobserved ,a tleas twithi non e vegetation type.Thre e species showed this feature, Lactarius chrysorheus 78 (fig. 24),Marasmius epiphylloides (fig.30) ,an d Typhula phacorrhiza (fig. 31). The clustering of M. epiphylloides ando fî \ phacorrhiza iscertainly - caused by the inhomogeneityo fth eplot s forthes especie s (seep .87) .I t was not possible to determine whether theplot s were heterogeneous for L. chrysorheus, nor to determinewhethe ri twa son ever yextende dmyceliu m (250-500m 2) pe rplot ,o r several smallermycelia ,growin gclustered .Th e clustering can eitherb ea characteristi c of L. chrysorheus, orca nb eth e resulto finhomogeneou splots . Legendt oth edistributio npattern s (Fig. 17-35). Abundanceo ffung ii sindicate da sfollows : PI onespecime npe rpartia lplo t I"! 2specimen s perpartia l plot [vl 3specimen spe rpartia l plot 171 4-10specimen spe rpartia l plot Ixl 11-29specimen s perpartia l plot C5 30-89specimen spe rpartia lplo t QQ 90-189specimen spe rpartia lplo t • >190specimen spe rpartia lplo t r indicates that the distribution was random, ctha tth edistributio nwa s clustered.Th ehighe rplan tcove ri sgive ni nBraun-Blanque tsymbols . —5> - o— -|-~ g- c e/ plot 12 plot 14 Fig. 17. Distribution and abundance of Clitocybe flaccida. 79 o e O i ooo o o O O O O O o o O O O O o ooo o o o o o o o o o o o o o o o o o o ooo plot 3 plot 4 plot 5 o ooo o o o o o o o o o 0 0 o o o "ö plot 9 plot 10 plot 11 plot 12 O 0 ooo o o o o o o o o plot 13 plot H plot 15 plot 20 Fig.18 .Distributio n of Scleroderma citrinum insevera lplots .Onl ypres encei sindicated . V • .. X . •. / / • 1 plot S plot 14 plot1 5 •• v c /— / / V •• .. .. /, • • :• . . V V V / • • c/ •• •• plot 16 plot1 7 plot2 0 Fig.19 .Distributio n andabundanc eo f Collybia peronata. 80 — _ • • c piot9 plot 22 Fig. 20. Distribution and abundance of Hapalopilus l-utilans. c • c c plot 10 plot 20 plot 22 Fig. 21.Distributio nan dabundanc eo f Tyromyces chioneus. • • / • c plotA plot 20 Fig. 22.Distributio nan dabundanc eo f Polyporus brumalis. 81 •• / / •• X c • • c c r plot9 plot 10 plot1 2 plot 20 Fig. 23. Distribution and abundance of Gymnopilus hybridus. , .. . • Xjv ••• • •• X]'• ' " X •• •• • >0V ••*/ • / c "•' • _ •• ij/ '•' " c •• • • plot 2 plot U, Fig.24 .Distributio n andabundanc eo f Lactarius chrysorheus. 7.3 CORRELATEDDISTRIBUTION S Inth eDicrano-Quercetum ,plot s 2 and 3, astron g correlationwa sob servedbetwee npartia lplot swit ha larg enumbe ro ffungu sspecie s (thati s morespecie stha nth emea nnumbe ro fspecie si nth epartia lplot so fa com positeplot )an dpartia lplot swit ha hig hcove ro fterrestria lmos s (cover >5%)(0.48* ,0.77*). M Three species showed asignifican t correlationwit hhig hmosscove r in bothplots : Marasmius androsaceus (0.59*,0.67*) , Galerina decipiens (0.61*, 0.34*), and Inocybe napipes (0.35*, 0.49*). Several other species hada significant correlation in one plot: Inocybe xanthomelas (0.51*, -), J. ovatocystis (0.41*, 0.28), Psathyrella fulvescens (0.36*, -), Collybia butyracea (0.35*, -0.17), Cordyceps ophioglossoides (0.32*, -), Galerina calyptrata (0.31, 0.46*),G . atkinsoniana (-, 0.52*), Nycena galopus (0.38*, 0.02), Russula emetica (-0.12,0.58*) , R. fragilis (0.08,0.56*) , Telephora terrestris (-0.12,0.45*) , Paxillus involutus (0.00, 0.40*), Xerocomus badius (0.15,0.43*) ,an d Amanita fulva (0.24,0.49*) .Se efig .2 5an d26 . 1) Values forV arementione d inparenthesis ,firs to fplo t2 ,secon dfo r plot3 .- mean sn oobservatio nbecaus eth especie sdi dno toccu ri ntha t ploto ri nles stha n3 partia lrelevés . 82 7 2 S 8 10 11 6 10 9 |12 2 ? 2 5 2 5 • • [ 9 9 3 12 9 IS 10 15 14 18 5 3 S 2 2 | 7 5 S S 10 16 14 13 IS 20 3 . 5 1S 1S 10 2 S 5 • /" ••• 9 6 4 14 14 16 8 9 26 18 20 10 10 50 4 15 5 number of species mosscover (terr.) in */. MarasmiuHit-Ss androsaceuE ZZs / . // V • • •• L V II X X « zzz -Jx 2KE m nocybe xanthomelas Galerinx/ a decipiens /P Cordycepsophioglossoide s I V I • • •'• / •• . . • '•" .. •• • nocybe napipes Inocybe ovatocystis Psathyrella"fulvescenI • s . .. / ...... •• • • • / • • Collybiabutyrace a Mycenagalopu s x/ Fig. 25.Distributio n of the number of species, of the moss cover, and the distribution and abundance of some fungi inplo t 2. Only very few species have a slightpreferenc e for growing in the spots with little moss. Few negative correlations of fungi with the moss patches were found, but only once was it significant: Phallus impudicus (-, -0.37*). The moss patches canb e considered as amicro-communit y with its own fun gus species. The moss patches were rather large, 25 to 300 m?, and covered 40% of the plot area inplo t 2 and 35%i nplo t 3. The moss patches indicate, and probably cause, theheterogeneit y of theD Qmentionne d onpag . 71.Whic h agents determine the growth of mosses is not known.A correlation with Quer- cus robur cover <75%exis t (0.29, 0.42), but if this is an agent it isprob ably not the only one. Partial plots with a high number of fungus species (more species than themea n number inpartia l plots in this composite plot) correlated signifi cantly with partial plots with a low cover of the herb layer (<10%) in plot 14. V=0.32*, see fig. 28.Partia l plots with a low cover of the herb layer are significantly clustered, as are partial plots with a high number of fungus species. Id o not know whether the fungi are the cause or the result of the absence of herbs, nor which factors cause the clustering. Lange (1923), Wilkins et al. (1938), Leischner-Siska (1939), Friedrich (1940), Kriegisteiner (1977) also reported that the richest fungus flora was found where phanerogam flora was poor. Few species correlated with a low cover of 83 u 7 #>• B 20 15 9 16 . • 15 IS 2Q 7 3 3 to IS 16 21 16 17 Tl 14 • 7 IS 7 5 7 2 Si 19 18 24 18 H S 1210 2 7 7 5 2 • • ZO 19 15 10 U 13 12 8 10 4 17 15 12 10 U 10 12 7 5 • • 2 2 • • number of species mosscove r (terr.)i n•/ . Marasmius androsaceus • : « 1 •/. . . •• • /// •• // . / / •••• • // • / .. • / •; • •• • / • / • . • • • • • • / • Inocybe napipes Galercno decipiens Galerina calyptrata ... v ...... • zz• • ^zz -zzz^ . /Z •• V Galerinz a atkinsonian• a -Russul • Ja emetica Russula fragilis Telephora terrestris Amanita fulvo Xerocomus badius . . / . . .• . / .. / / .• / / / / V •• / / •• " V •• /^ •• •• Paxillus invoiutus Phallus impudicus Fig.26 .Distributio n ofth enumbe ro fspecies ,o fth emos scover ,an dth e distributionan dabundanc eo fsom efung ii nplo t3 . theher b layer (fig.27 ) Clitocybe vibecina V=0.49*an d Oudemansiella pla- typhylla V=0.43*; and some other specieswit ha lowe rvalu efo rV an dno t significant such as: Calocera cornea, Stereum rugosum, Pluteus cervinus, Rutstroemia firma, Marasmiellus ramealis, Phlebia radiata, Crepidotus pubes- cens, Collybia peronata. These are all species growing on litter (fallen leaveso rtwigs )o ro nfallen ,o rerect ,dea dwood . High density of theher b layer inplo t1 4wa scause db yhig hdensitie s of Maianthemum bifolium, Deschampsia flexuosa, and Hedera helix, onlyo ri n combination, and never exceeded40 %o f apartia l relevé. Marasmius epi phylloides (V=0.91*,se efig .30 )an d Mycena cinerella (V=0.63*)correlate d with Hedera helix. Noothe rfung iappeare dt ocorrelat ewit ha hig hdensit y ofth eher blaye ro rwit ha specifi cherb . 84 10 7 5 10 7 10 7 7 . • .. .. - •• 7 20 20 S 7 7 7 10 • • • •• 10 10 5 2 7 10 25 30 10 7 7 5 7 10 30 40 // H IS 10 IS 10 20 30 30 30 • / •• mAAAA-\/V\ cover of herb layer in•/ . Deschompsiaflexuos a Maianthemum bifolium 26 13 222 42 1 2619 •• / * XXX .. . .. 20 21 23 21 / /•• / V .. • • XXX 7 222 6 / / • / • / • 22 •• / X /x / number of fungus Ctitocybevibecina Oudemansiella piatyphylla species OO O O ° V z.. ..• O O o 2» ° o Crepidotus pubescens Rutstroemia firma Stereum rugosum • / / •• § Mycena cinerella Laccaria proxima Mycena inclinata / / .. s?ï= • [2 • Hypholccn•a fascicular e Pluteus cervinus Psathyrella squamosa r I I 1^ Fig.27 .Distributio no fth eher blaye rcover ,o fth enumbe ro ffungu sspe cies, and the distributionan dabundanc eo fsom eplan tan dfungu s speciesi nplo t14 .(I n Rutstroemia firma and Stereum rugosum only- presencewa sindicated. ) 85 cover of the herb layer in •/. 40- 30' 20^ •• • • • KA- 20 25 number of fungus species Fig28 . Relation between thenumbe r of fungus speciesan dth eher blaye r coveri nth epartia lrelevé so fplo t14 . 86 r 2b 3 2b 2b 2b 2b r 2a r 2rr 1 + 2b 3 3 3 3 3 2b 2a + 1 1 2b Quercusrobu r Betuia pubescens Vaccinium myrtillus Ilex aquifolium . Z•• • ^ 0 o • ^Z 0 ZZ^ _ i 0 o ÈZZ444"Z lactarius quietus Paxillus involutus Myceno vitilis I "~' X • • • I Panellus serotinus Pkit e us C(>r \ in us Collybia butyracea 10 10 11 7 9 9 9 10 12 9 10 9 12 11 •n 8 10 12 10 12 11 a 9 9 10 10 10 10 10 9 9 6 • • 14 12 11 11 7 9 11 11 Mycena einerella Collybia peronata number of species inth e partial plots Fig.29 .Th e distribution and abundance of somehighe rplant s andfungi , andth edistributio no fth enumbe ro ffungu sspecie si nplo t24 . (In Paxillus involutus onlypresenc ewa sindicated. ) Inplo tnumbe r2 4th eplan tspecie s Quercus robur, Betula pubescens, and Vaccinium myrtillus wereno thomgeneousl ydistribute dbu tgre wtogethe ri n only a smallpar to fth eplo t (fig.29) .Th edensit yo f Ilex aquifolium in thispar twa s slightly less than inth eres to fth eplo t (fig.31) .Shru b andher blaye rwer ealmos tabsen ti nth ewhol eplot . 87 Several fungus species followed Quercus robur, Betula pubescens, and Vaccinium myrtillus. Correlationswer e (valuesfo rV ar egiven) : with Q. robur B. pubescens V. myrtillus Lactarius quietus 0.73* 0.60* 0.65* Paxillus involutus 0.26* 0.48* 0.22 Mycena vit il is 0.51* 0.24 0.43* id.^ 4specimen s 0.91* 0.49* 0.83* Panellus serotinus 0.22 0.40* 0.20 Pluteus cervinu s 0.38* 0.28 0.35* Collybia butyracea 0.38* 0.28 0.35* C. peronata 0.38* 0.28 0.35* Mycena cinerella 0.38* 0.28 0.35* Correlationbetwee n the fungus speciesmutuall y wasver ylo wan dno tsig nificant.Thes e fungiprobabl y depended onthes eplants .Th einfluenc eo f theplant swa sno tver y great.Th enumbe r of fungusspecie si nthi spar t wasno tlarge rtha ni nth eothe rpart so fthi splot . The 2 other Violo-Quercetum ilicetosum plots were morehomogeneou s; especially Q. robur was homogeneously distributed over the whole plot (SF100%) , son o correlationswit hfung icoul db ecalculated .No rcorrela tionswer e found ofthes efung iwit h B. pubescens norwit h V. myrtillus in theseplots . Heterogeneity of aplo t is also shownfo rfung idependin go na specia l substrate,suc ha s (litter of)a specificplant ,o r anotherfungus .Thes e substrates are often not quitehomogeneousl y distributed throughoutth e plot, so theplo t isno thomogeneou s forthos e fungi.Som e examples are discussedbelow . Thesubstrat efo r Marasmius epiphylloides isdea dleave so f Hedera helix. Thefungu swa sfoun di n3 plot san dha da correlatio nwit h H. helix growing terrestrially, cover >5%o f 0.91* inplo t14 ,o f0.46 *i nplo t15 ,an do f 0.43* inplo t1 7 (fig.30) .H. helix hada clustere ddistributio ni nthes e 3plots ,s oth eplot swer einhomogeneou sfo r M. epiphylloides. Asa resul t the distribution of M. epiphylloides was also clustered. Iti sno tknow n which agentsprevente d M. epiphyloides from growing inal lpartia l plots witha 5 %o rmor ecove ro fterrestria l H. helix. The leaves of Ilex aquifolium areth e substratefo r Typhula erythropus and T. phacorrhiza. These fungiwer e only found inth eplot s2 0 and24 , where they correlated with I. aquifolium in the tree layer cover>75 % (fig.31) .Value sfo rV arefo r T. erythropus 0.56*i nplo t2 0an d0.63 *i n plot24 ,fo r T. phacorrhiza 0.70* inplo t2 0an d 0.45*i nplo t24 .Appar ently the Ilex litter is only suitable for the Typhula species ifth e density ofth e Ilex treesi shigh .Th efung iwer eno tfoun di nothe rplot s withhig h Ilex densities.Apparentl yther ear eothe rfactor stha tprevente d thefung ifro mgrowin gthere . 88 I 1 2fr2m 1 3 3 2b 2t + r + 2m 2a 2b 2b 2b 3 3 1 2rr + + + + + l + 2m 2a 2a 3 2b 3 3 3 2a 1 + 1 + + + + + + 2rr 2mCoCo 1 2ir 2a 2a 3 3 2b 2b 2b 2b 2b Zb Zb Zb 2b 2b 2b 2b 2a 3 [2b Pb 2m 2rr2m2o 3 4 4 3 //* OlOIO :Ä plot 15 pbtK plot 17 Fig.30 .Distributio nan dabundanc eo fterrestria l Hedera helix (above)an d of Marasmius epiphylloides (below,i nplo t1 5onl ypresenc ei s indicated). 2b 2b 2b . plot 20 Zb 2t S . / 2b 3 2b S S S 5 4 S 5 5 / S S S S 5 5 5 4 plot 24 s U 5 S 5 S S t m 4 2b 5 4 4 S S S w • 3 2 2 t. 4 5 S 4 fi$+ ••• •• Ilex aquifolium Typhula phacorrhiza Typhula erythropus Fig.31 .Distributio nan dabundanc eo f Ilex aquifolium (treelayer) , Typhula phacorrhiza, and T. erythropus inplo t2 0an d24 . 3 2b + + r 3 4 3 2b 4 2a . 4 2b 3 2b + + 3 . 2b 3 + + 4 4 . . 2b 3 + + + + 3 4 • Pteridium aquilinum Psilocybe crobula Fig.32 .Distributio nan dabundanc eo f Pteridium aquilinum ando f Psilocybe crobula inplo t20 . 89 Psilocybe crobula was foundt ogro wo nth epetiolu so fleave so f Pteri- dium aquilinum in plot2 0 (fig.32) .Correlatio n was 0.52*.Correlatio n with P. aquilinum cover >50%wa smuc hhigher ,0.90* .Apprentl y P. crobula prefers high densities of P. aquilinum. The speciesals ogre wtogethe ri n two otherplots ,bu tn ocorrelatio nwa sfound ,-0.0 8i nplo t15 ,an d 0.07 inplo t17 . / • / •; x^XSZ z Armillariella *—rJK— mellea v •: . SZZSSz§/ !z V • • m KB Collybia s • V cookei m EEK ' •• 5 m *§* *>X/ plot 5 plot1 2 plot U Fig. 33.Distributio n and abundance of Armillariella mellea and Collybia cookei inplo t5 ,1 2an d14 . . £S ••g zg^ • EX X& x wàâ Q z zzs coo ei • • -y— plot2 0 plot2 2 Fig. 34 Distribution and abundance of Hypholoma fasciculare and Collybia cookei inplo t2 0an d22 . 90 Collybia cookei growso n 'rotten'flesh yfungi .I nfac tth esubstrat ei s not 'rotten',bu t it isa kin d of Sclerotium. Insevera l plot C. cookei grew on Armillariella mellea (fig.33) .I nplot swher e both species grew together, in at least2 and atth emos t all but 5partia l relevés,th e followingvalue s forV wer e found:plo t5 :0.55* ,plo t10 :0.25 ,plo t11 : -0.17, plot12 :0.35* , plot16 :0.22 ,plo t17 :0.18 ,an dplo t22 :0.05 .I n theplot s5 an d1 2th edistributio no f C. cookei depended,a tleas tpartly , onth e distribution ofA . mellea. Inplo t1 4th eSpatia lFrequenc yo fbot h specieswa s very high, 98%,s o acalculate d correlationwa smeaningless . The substrate for C. cookei was distributed homogeneously as alsowa sC . cookei. Iti sclea r that also in thisplo t thedistributio no f C. cookei depended verymuc h on thato fA . mellea. Otherspecie so nwhic h C. cookei could depend, like Paxillus involutus, Russula or Boletus species,d ono t explainth edistributio n ofC . cookei inth eplot s5 ,12 ,an d14 .I nth e plots2 0 and 22 thedistributio no f C. cookei wascorrelate dwit hth edis tributiono f Hypholoma fasciculare, V=0.57*,an d0.38* . (fig.34) .I nothe r plots where these species grew together no significant correlationwa s found. Paxillus involutus is amycorrhiza lsymbiont .I nsevera lplot sa stron g correlation with Betula pubescens cover>12.5 %and/o rcove r>25 %wa sfoun d (fig.35) .Th e followingvalue s forV wer efoun d (onlycalculate d inplot s were P. involutus had a spatial frequency ofa tleas t5 %an d B. pubescens atth emos ta spatia l frequencyo f 90%): correlation(V) of Paxillus involutus SpatialFrequenc yo f with Betula pubescens Betula pubescens plotnr . cover >12.5% >25% cover >12.5% >25% 5 0.41* - 12 0 9 0.19 0.35* 88 50 10 0.17 0.26 71 35 11 - 0.13 93 43 13 0.25 0.41* 80 38 14 0.43* 0.31 35 32 15 0.13 0.14 80 50 16 0.14 0.09 53 8 24 0.46* 0.48* 20 8 The specieswer e significantly correlated inth e 3plot swit h thelowes t frequencyo f B. pubescens. In2 o fth e4 plot swit ha hig hfrequenc y (>80%) of B. pubescens therewa s acorrelatio nwit h B. pubescens cover >25%.N o (significant)correlatio nwa s foundi n4 plots . P. involutus canals ofor m mycorrhizawit h Quercus robur. Thiswa sapparentl yth ecas ei nth eplot s2 , 3, and 4,wher e B. pubescens did notoccur .N ocorrelatio nwa s foundwit h otherhost-plants ,suc ha s Populus tremula or Fagus sylvatica. 91 So • 2o plot 5 . 2b 2b . . •• • .. 2o • • • • • •• 2b 3 2o 2b 3 2b 2o 2b 3 2a • 2b 3 3 2o 4 4 2o 3 2a plot 9 • • / •• 3 Zb 5 2b 3 3 3 4 3 • • 4 21 3 3 4 4 2b 2b • + 2b 2a 2b 2a 2a 2b 2b 3 + 2b 2a 2a . plot1 3 2b 3 3 4 2b 3 .. 3 5 3 2b 2a 3 2b 2a 3 3 3 5 3 4 2c • / 3 2b 4 3 • 3 + 2b 2b • plot U 2a • 3 3 3 4 2b 3 • r plot 24 2b 3 • • 2b r 2a 3 3 2b 2a •• Betula pubescens Paxillus involutus Fig.35 .Distributio n and abundance of Betula pubescens intre elayer ,an d Paxillus involutus insevera lplots . Between Paxillus involutus and Phallus impudicus anegativ ecorrelatio n was found in severalplots .Th e followingvalue s forV were found (only calculated in plots where both species occurred at least in2 partia l plots):i nplo t3 :-0.59* ,i nplo t5 :-0.02 ,i nplo t11 :-0.21 ,i nplo t13 : -0.14, inplo t15 :-0.05 ,i nplo t16 :-0.14 . Inplo t3 ,th eonl yon ewher e thecorrelatio nwa s significant, P. involutus grewi nth emos spatche san d P. impudicus outsideth emos spatche si nth ebar elitte r (fig. 26). 92 8 COMPOSITION OF ECOLOGICAL GROUPS Withinth e fungusvegetatio ni nthes eoakwood s6 ecologica lgroup swer e distinguished: mycorrhizal species;saprophyte s onhumus ;saprophyte s on decaying wood and branches; saprophytes on fallenleaves ,fruits ,dea d herbs, andmosses ;parasites ;an d fungi on dung andbir d pellets.Ever y specieswa s classed inon eo fthos egroups .Som especie swer edifficul tt o class, e.g. it isno t always knownwhethe r aspecie si smycorrhizal .Fo r the species ofgener a such as Cortinarius and Lactarius thisquestio ni s beyond doubt,bu t ingener asuc ha s Inocybe and Tricholoma opinionsar ea t variance. Species can also be optionally mycorrhizal,suc h as Laccaria species.Followin gliteratur e (mainlyTrapp e1962 )I considere d asmycorrhi zal the species ofth egener a Amanita, Boletus, Cantharellus, Cortinarius, Dermocybe, Hebeloma, Hydnellum, Inocybe, Lactarius, Leccinum, Paxillus, Phallus, Phellodon, Phylloprus, Russula, Sarcodon, Scleroderma, Tricholoma, Tylopilus, Xerocomus (except X. parasiticus), and Lepista nuda. The saprophytus were splitu p in3 groups ,terrestria l growing humus saprophytus,saprophyte so ndecayin gwoo do fstumps ,trunks ,logs ,branches , and saprophytes on litter, fallen leaves,fruits ,dea dherbaceou splants , andmosse s (shortly calledlitte rsaprophytes) .Saprophyte sgrowin go nde cayingwoo do fstumps ,trunks ,logs ,an dbranche sar egenerall yeas yt odis tinguish,a sthi swa yo fgrowin gi susuall ya clea rcharacteristic .Classe d aswoo dsaprophyte sar eth especie so fth e Polyporaceae, ofth egener a Bul garia, Calocera, Coryne, Chondrostereum, Exidia, Gymnopilus, Hohenbuehelia, Hypholoma, Kuehneromyces, Lentinellus, Marasmiellus, Nectria, Oudemansiella, Panellus, Pleurotus, Pluteus, Rutstroemia, Sphaerobolus, Steccherinum, Tremella, Tricholomopsis, Xylaria, and Collybia fusipes, Coprinus sect. Micacei, Crepidotus haustellaris, Entoloma euchrous, Galerina cinctula, G. ampullaceocystis, G. triscopa, Mycena galericulata, M. haematopus, M. inclinata, M. polygramma, Pholiota alnicola, P. tuberculosa, Psathyrella hydrophila. Itwa s not always easy to distinguish thehumu s saprophytes fromth e littersaprophytes .I twa sdifficul tdetermin ewhethe rsom especie sgro wt o thehumu s or onth efalle nleaves .I considere dhumu ssaprophyte sth espe cies ofth e genera Clavulina, Clitocybe, Cystoderma, Entoloma (except E. euchrous), Helvella, Laccaria, Lepiota, Psathyrella (exceptP . hydrophila), Tephrocybe, Tubaria, and someo fth especie so f Collybia, Mycena, Stropha- ria. Inth egrou p of litter saprophytes itwa s sometimes difficultt odis tinguish between saprophytes and parasites. Especially in some species 93 growing onherbs ,suc h as Mycena rorida, Crepidotus pubescens, itwa sno t clear whether they grew on dead or on still livingherbs ,o ri nothe r words, whether they were saprophytes orparasites . Iconsidere d thema s saprophytesjus tlik e Collybia cirrhata, C. cookei and C. tuberosa. Species (onlya few )growin go no rbetwee nmosse swer eals oadde dt othi secologica l group.I considere d aslitte rsaprophyte sth especie so fth egener a Ciboria, Clavariadelphus, Collybia (exceptC . fusipes), Crepidotus (except C. haus- tellaris), Marasmius, Psilocybe, Rickenella, Typhula, andsom eo fth especie s of Galerina, Mycena. The ecological group ofth eparasite s hadonl ya fe wspecies .Th edif ferencebetwee nparasite s and saprophytes isno talway swel ldefined .Tw o species were included thatar esometime sparasite s and sometimessapro phytes: Armillariella meJlea and Heterobasidion annosum. Parasiteso nin sects and other fungi were alsoinclude dher e (Cordyceps species).Th e group of fungio n dung andbir dpellet si sals over ysmall :som e Coprinus species, Onygena corvina, Panaeolus fimicola, Stropharia semiglobata. Thenumbe r of speciesi neac hecologica lgrou pwer ecounte dpe rvegeta tion type and foral lvegetatio ntype stogethe r (table 10).Th eecologica l group of themycorrhiz a species isespeciall y important inth eDicrano - Quercetum.Man y ofth emycorrhiz aspecie sfoun di nthi sstud ywer epresen t here,an dnearl yon ethir do fth enumbe ro fspecie si nth eDicrano-Quercetu m weremycorrhiz aspecies .Th enumbe ran dproportio no fmycorrhiz aspecie swa s muchlowe ri nth eQuerco-Betuletum , andstil llowe ri nth eViolo-Quercetum . The groupo fth ehumu s saprophytes isalmos tequall y importanti nal l the vegetations types. Thenumbe r inth eViolo-Quercetu m ilicetosumwa s slightly lower,bu t inproportio n itequale dtha ti nth eothe rvegetatio n types. The group ofsaprophyte s onwoo d isespeciall yimportan ti nth eViolo - Quercetumilicetosum ,wher e6 2specie swer epresent .Nearl yhal fth enumbe r of species in this sub-association werewoo d saprophytes.Abou tth esam e numbero fspecie swa spresen ti nth eViolo-Quercetu mtypicu man dth eQuerco - Betuletum, butther e itwer e smallerproportion s ofth e totalnumbe r of species.A muc hsmalle rnumbe r (andproportion )wa spresen ti nth eDicrano - Quercetum. The lownumbe r inth eDicrano-Quercetu m isprobabl y causedb y the sparse covering ofth emicrohabita t of 'dead stumps andlogs' ,0.07 % (seetabl e 2). Thismicrohabita t ismor e frequenti nth eQuercio nrobori - petraeae,coverin g1.5 %i nth eQuerco-Betuletum , 1.6%i nth eViolo-Quercetu m typicum,an d1.0 %i nth eViolo-Quercetu m ilicetosum. The ecological group ofsaprophyte so nlitte ri slarges ti nth eQuerco - Betuletum and theViolo-Quercetu m typicum.A smalle rnumbe rwa spresen ti n theViolo-Quercetu m ilicetosum,bu ti nproportio n itequale d that inth e Querco-Betuletum and theViolo-Quercetu m typicum. Inth eDicrano-Quercetu m thenumbe r (andproportion )wa sslightl ylowe rtha ni nth eassociation so f theQuercio nrobori-pertraeae . 94 Table10 :Ecologica l groups.Numbe r ofspecie si nth e6 group spe rvegeta tiontyp e (inparentheses :percentag eo fth etota lnumbe ro fspe ciesi ntha tvegetatio ntype )an do fal lth evegetatio ntype sto gether.M=mycorrhiz a species,H=humu ssaprophytes ,W=saprophyte s ondecayin g wood, L=saprophyteso n litter,an dbranches ,falle n twigs, leaves or fruits, dead herbaceous plants andmosses , P=parasites,D=fung io ndun go rbir dpellets . M H W L P D Dicrano-Quercetum 78 (44) 34 (19) 36 (20) 23 (13) 4 (2) 4 (2) Querco-Betuletum 40 (21) 41 (22) 56 (30) 41 (22) 5(3 ) 4 (2) Violo-Quercetum 2Q (lfi) 4Q (M) 6Q (34) 3g (M) 3 (2) g (3) Violo-Quercetum^ lg (14) 25 (lfl) 62 (4fi) 2? (2Q) ± (1) 2 (1) total 102 (33) 64(20 ) 81 (26) 52 (17) 7(2 ) 7(2 ) The groups ofth eparasite s and ofth especie so ndun gha donl ya ver y small share inth ecompositio no fth emycoflora ,whic hdi dno tdiffe rmuc h inth e4 (sub-)associaions . The Querco-Betuletum had about the same share of saprophytes, but slightly moremycorrhiz a speciestha nth eViolo-Quercetu m typicum.Th elo w numbero fspecie si nth eViolo-Quercetu m ilicetosum ismainl yth eresul to f thever ylo wnumbe ro fmycorrhiz aspecies ,an dth eslightl y lowernumbe ro f humus and litter saprophytes.Th eDicrano-Quercetu m hadles ssaprophytes , but many more mycorrhiza species thanth e associations ofth eQuercio n robori-petraeae. 95 SUMMARY In this thesis the fungusvegetation , thevegetatio n ofphanerogams , mosses, and lichens, and the soil (profiles and chemical analysis)ar e described in the associations Dicrano-Quercetum, Querco-Betuletum, and Violo-Quercetum, both typicum and ilicetosum inth eprovinc eo fDrenth e (NorthEas tNetherlands) . Thevegetatio n ofphanerogams ,mosses ,an d lichenswer edescribe dwit h the aid ofrelevé saccordin gt oth eBraun-Blanque tmetho dwit ha modifica tion according to Barkmane t al. The synusiao fmosse s ondea dtruncs , stubs, and logswa s separately described,becaus ei ti sth esubstrat efo r somefungu sspecies .Th ecorrec tname so fth esyntax aan dth esynsystemati - calpositio nwer e determinedwit hth eai do fliterature .Th echaracte ran d differential speciesmentione d in literaturewer e discussed.Th eQuerco - Betuletum and theViolo-Quercetu m appeared tob eclosel yrelate dassocia tions,belongin g toth eallianc eQuercio nrobori-petraeae .Th eViolo-Quer cetumilicetosu mwa sdescribe da sa ne wsub-association .Barkman' sopinion , the Dicrano-Quercetum is anindependen t association,wa s confirmed.Thi s associationha sman ydifferentia lspecie s (almostonl ymosse san dlichens) , thereforeth eaffinit ywit hth eassociation so fth eQuercio nrobori-petraea e islow ,an di ti sno tinclude di nthi salliance . Theplot so fth eQuercio nassociation swer eal lsituate do nsand ysoils , oftenwit h asubsoi l ofloa m or loamy sand.On eplo tha da 'broek'eart h soil, the other had distincthumu s podzolprofiles .Th e Violo-Quercetum ilicetosumplot sha drelativel ydee pprofiles ,wit hofte na thic kAl hlayer . Inth eViolo-Quercetu m typicum and theQuerco-Betuletu m theprofile swer e moreshallow ,an da nAl hlaye rwa susuall yabsent .Th etopsoi li sth eQuer co-Betuletum plots was generally disturbed. TheDicrano-Quercetu m plots appeared tob e located on soils ofyoun g drift sands,i nwhic ha profil e wasonl yweakl ydevelope d ('duin',vagu esoi lwit ha micropodzol) . Chemical analyses weremad e from samples ofth eA Oan dth eA2 1layers . Inthes elayer sth elarges tamoun to fmyceli ai spresent .Th esoil sappeare d tob e very acid,p H2. 9 -3. 5 inth eA2 1layer ,3. 3 -4. 7 inth eA Olayer . Inth eA Olaye rNa ,K ,Mg ,an dC aion s (giveni nmas sfractio nan dmas scon centration)wer epresen ti nalmos tth esam equantitie si nal lth edifferen t types ofvegetation . Inth eA2 1laye rth e concentrationswer e highesti n the Violo-Quercetum ilicetosum, and lowest inth eDicrano-Quercetum . The 'total ion supply'wa shighes ti nth eViolo-Quercetu m ilicetosum,wa smuc h lower inth eViolo-Quercetu m typicum and theQuerco-Betuletum , andlowes t inth eDicrano-Quercetum .Th eC/ Nratio ,however ,increase d inthi ssequence . 96 Description of the fungusvegetatio nwa sbase dupo nsynthetica lrelevé s ofpermanen tsampl eplot sstudie ddurin gfou ryear si nsuccession .Th enum ber of fungus species (314)largel y excededtha to fphanerogam san dfern s (72)o rmosse san dlichen s (67).Criteri awer edraw nu pt odistinguis hdif ferentialspecie si nth efungu svegetation . TheDicrano-Quercetu m and theQuercio nrobori-petraea ehav ei nrelatio n toeac hothe rman y differential species.Thi s supports the ideatha tth e Dicrano-Quercetum does not belong to the Quercion robori-petraeae.Th e Querco-Betuletum and the Violo-Quercetum have in relation toeac hothe r less differential species,bu t are also good characterized. TheDicrano - Quercetum and theViolo-Quercetu m ilicetosum, bothver ypoo r invascula r plant species,appeare d withth eai do ffungu sspecie st ob echaracterize d better.Thi s holds especially forth eViolo-Quercetu m ilicetosum,whic hi s alsover ypoo ri n(terrestrial )mosse san dlichens . The 'minimalarea' ,o ractuall yth erepresentativ eplo tsiz eappeare dt o be 1500m 2 inth eQuercio nassociations ,an dt ob ea tleas t300 0m 2 inth e Dicrano-Quercetum. A new characteristic, the 'Total Spatial Frequency' (TSF)ha s been created to give an aproximation ofth e real abundance offungi ,i.e .th e abundance ofmycelia :th eproportio n ofth enumbe r ofpartia lplot so fa permanent plot in which carpophores oftha t specieswer e ever observed duringth eresearc hperiod .Th eabsolut emaximu mabundanc e (AMAC)doe shav e acertai ncorrelatio nt oth eTSF ,bu ti ti sgenerall yimpossibl et oconver t onecharacteristi c intoth eother . Thedeterminatio no fth eTS Fals oyielde dresult so nsom eothe raspects . Thus itwa spossibl et oprov etha tRaunkiaer sla wo fconstan tspecie sals o applies to the fungio fth eQuercio nrobori-petraeae ,bu tno tt oth efung i ofth eDicrano-Quercetu m (incontras tt oth evegetatio no fgree nplant so f theDicrano-Quercetum) .Th eDicrano-Quercetu m appeared tob echaracteris - ticaly heterogeneous,mor e heterogeneoustha nth estand so fth eothe rsyn - taxa,owin gprobabl y to the largemos spatches .Distributio npattern so f thespecie swithi nth eplot swer eals ostudied .Th edistributio nwa smostl y random, and never underdispersed (regular).Onl y 35fungu s specieswer e clustered (overdispersed) intw oo rmor eplots .Thi sma yeithe roccu ri fa mycelium is soextensiv etha ti ti ssprea dove rtw oo rmor epartia lplots , or if the mycelia were clustered, e.g. because of inhomogeneity ofth e plot,a tleas twit hregar dt oenvironmenta l factorsaffectin gthi sspecies . Someexample swer estudie do fcorrelate ddistribution so ffung ii nrelatio n toeac hother ,o ro ffung ii nrelatio nt ovascula rplant so rmosses .Severa l fungus species (e.g. Marasmius androsaceus, Galerina decipiens, Inocybe napipes) had acorrelatio nwit hth emos spatche si nbot hth eDicrano-Quer cetumplots .I n aViolo-Quercetu m typicumplo thig hnumber so ffungu sspe ciesi npartia lplot scorrelate dwit ha lo wcove ro fth eher blayer . Maras mius epiphylloides correlated with terrestrial Hedera helix, butdi tno t 97 occur in allpartia l plots with ahig hcove ro f H. helix. Collybia cookei correlatedwit h Armillariella mellea insom eplots .Som eothe rcorrelation s werediscussed . The fungiwer e classed inecologica l groups (themai ngroup swer eth e mycorrhizal species,th e saprophyteso nhumus ,th esaprophyte so nwoo dan d branches,an dth esaprophyte so nleaves ,fruits ,herbs ,an dmosses )an dth e share of each ofth e groups inth e fungusvegetatio nwa sdetermined .Th e Dicrano-Quercetumappeare dt ohav ever ymany ,th eViolo-Quercetu milicetosu m very fewmycorrhiza l species.Th eDicrano-Quercetu m hadth efewes tnumbe r ofsaprophyte so nwood ,th eViolo-Quercetu m ilicetosum thefewes tnumbe ro f humus saprophytes.Th e numbero fsaprophyte so nleave setc .wa shighes ti n theQuerco-Betuletu m andth eViolo-Quercetu mtypicum . Shorto r longernotes ,o r descriptions aregive no fa numbe ro ffungu s species with indentification difficulties,cause d by themateria l orb y taxonomical problems.A newfungu staxo ni sdescribed : Psathr/rella fulves- cens var. dicrani. 98 LITERATURE Bakker,H . de,& J.Schelling , 1966.Systee m vanbodemclassificati e voor Nederland.Wageningen . 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Ypelaar,P. ,i nprep .Mycosociologisc honderzoe kva nhe tDicrano-Quercetum . 103 LIST OF PLOTS APPENDIX 1 Foreac hplo ti sgiven :Provinc e (Dr.=provinceo fDrenthe ,Ov.=provinc e ofOverijssel) ,Municipality ,an dmor edetaile dindicatio nsuc ha sth enam e ofth e region;Tm=Topographica lmap , followed byth enumbe ran dlette ro f thesheet s1:25,000 ,an dth ema pcoordinates ;th esampl eplo tarea ,an d(i f itconcerne da composit eplot )th eare aoutsid eth eplot ,i nth esam easso ciation,tha twa s alsoinvestigated ;a shor tcharacterizatio no fth evege tationan dth egrowt hfor mo fth ewood ,an dsometime sparticular so fag eo r situation; situation inth e form-units ofth eGeomorphologica l map sheet 17+18, (Stiboka 1978)- a sn o other sheetswer eavailabl eform-unit swer e onlygive nfo rplot slyin go nthi ssheet .Ol dtopographica lmap swer estud ied to determine how old thewood swere .Especiall y the series1:25,00 0 reconnaissance between 1896-1900 (published 1902-1904),bu tth e slightly older map 1:50,000, and the "Choro-topographische Kaartde rNoordelijk e Provinciënva nhe tKoninkrij kde rNederlanden" ,scal e1:115,200 ,publishe d 1823 (facsimilereprin tHaarle m1979 )wa sals oconsulted . 1.Dr. ,Beilen ,Boswachteri jDwingelo oplo t59 ;T m17C ,536. 9- 226. 6Are a 800m 2. Schrubby oakwoo d almostwithou t aher blayer ,bu twit hextensiv e localmos s 'carpets'-Lyin go na lo winlan dsanddun e (4L8).Probabl yorigi natedo rplante dabou t1920 . 2.Dr. ,Beilen ,5 k mN Wo fth evillag eo fBeilen ,Zui dHijkerzand ;T m17A , 546.7- 227.3 .Are a100 0m 2 +abou t100 0m 2. Scrubbyoa kwoo dalmos twithou t aher blayer ,wit hextensiv eloca lmos s 'carpets',surrounde db yPin ewood s andheathlands .Situate d ina nare ao flo winlan dsanddune san dassociate d plains (4L8). Thewood sprobabl y originated orwer eplante d between185 2 and1896 . 3.Ov. ,Steenwijk ,5 k mN o fth evillag eo fSteenwijk ,Heerlijkheid "d eEese ; Tm16E , 538.1- 203.7 . Area 1000m 2 +abou t200 0m 2. Oak coppicewit ha sparseher blaye rwher e Deschampsia flexuosa dominated,wit hextensiv eloca l moss 'carpets'.Probabl y lying in an area of inland sanddunes.Th ewoo d originatedo rwa splante dbefor e1932 . 4.Dr. , Rolde, 1k mN ofth evillag e ofSchoonloo ,Schoonloë rstrubben ; Tm17E , 547.9- 243.2 . Area100 0m 2 +abou t 500m2 . Scrubby oak-birch woods bordering on heathland, with a herb layer ofmainl y Vaccininium myrtillys, V. vitis-idaea, and Calluna vulgaris. Situated inlo winlan d sandduneswit h associated plains (4L8). Thewood soriginate dbetwee n185 2 and189 6fro mspontaneou sregeneratio no fth ewood so nheathland . 104 5.Dr. , Ruinen, 3k mE of Pesse, Amshoff's bos; Tm17C ,531. 7- 229.9 . Area105 0m 2 +abou t100 0m 2. Oak-birchwoo dwit h aspars eher b layero f Vacciaium myrtillus. "Spaartelgen"woo d (former coppice,transforme dint o highwood) . Situatedi na relativel yhig hare ao fgroun dmorain ewit hcover - sand (3L2a).Th ewoo dwa splante dbetwee n185 2an d189 6o nheathland . 6.Dr. ,Ruinen , 1k mN ofPesse ;T m17C ,533. 4- 226.7 .Are a150 0m 2.Hig h oak-birch wood, theher b layerwit h agrea t deal of Vaccinium myrtillus. Situatedi na relativel yhig hare ao fgroun dmorain ewit hcoversan d (3L2) . Thewood s wereplante dbetwee n185 2an d189 6o nheathland ,probabl yorigi nallywit hconifers ,whic hwer elate rreplace db yoaks . 7.Dr. ,Oosterhesselen ,Havezath e DeKlencke ;T m17G , 531.7- 246.2 .Are a 2000m 2. High oak-birch wood with agrea t deal of Vaccinium myrtillus. Situatedi na relativel yhig hare ao fgroun dmorain ewit hcoversan d (3L2a). Thiswoo d already existedi n1852 ,probabl ythe nwit hconifers ,whic hwer e laterreplace db yoaks . 8.Dr. ,Norg ,2 k mS o fth evillag e ofNorg ,Tonckensbos ;T m12A ,562. 6- 226.1. Area200 0m 2. High oak-birchwoo d with agrea t deal of Vacciaium myrtillus and Helampyrum pratense. An oldwood ,alread ymarke do nth ema p of1823 . 9.Dr. ,Westerbork , 1k mNN W of the village of Mantinge,Mantingerbos ; Tm17D , 536.5- 236.9 . Area100 0m 2 +abou t100 0m 2. Oak-birch woodwit h a greatdea l ofVacciniuj n myrtillus. Coppice,whic hha sno trecentl ybee n cut, situated on araise d ground moraine in abroo kvalley .A nol dwood , alreadymarke do nth ema po f1823 . 10.Dr. ,Vries ,1 k mN W of thevillag e ofZeijen ,Zeijerstrubben ;T m12B , 563.8- 232.2 . Area105 0m 2 +abou t50 0m2 . Oak-brich wood with a great deal of Deschampsia flexuosa, and some Vaccinium myrtillus and Molinia caerulea. Coppice,no t recently cut.Situate d ina nol dstri po fwoodlan d surroundingth ecommo nfields ,alread ymarke do nth ema po f1823 . 11.Dr. ,Wersterbork , 1k mNN W ofth evillag e ofMantinge ,Noordlagerbos ; Tm17D , 536.9- 237.4 . Area 1000m 2 +abou t100 0m 2. Oak-birch woodwit h a great dealo f Vaccinium myrtillus andsom e Pteridium aquilinum. "Spaar- telgen"wood , situated on arelativel yhig hplai n ofgroun dmorain ewit h coversand (2M5)i n abroo kvalley .Marke da swood so nth ema po f1823 ,a s cultivated lando nth e185 2map ,an dagai na swood so nth e190 4map . 12.Dr. , Beilen, Boswachterij Hooghalen, Heuvingerzand; Tm17B , 547.2- 233.1. Area105 0m 2 +abou t50 0m 2. Oak-brich wood with amongst others Lonicera periclymenum, Deschapsia flexuosa, and Maianthemum bifolium. Cop pice,no trecentl ycut .Situate d ina relativel yhig hare ao fgroun dmorain e with coversand (3L2a). Originatingbetwee n 1852 and 1896b yplantin go r spontaneous regeneration ofth ewood s onheathlands ,directl ybordere db y theplante dstri po fwoodlan dsurroundin gth ecommo nfield so fth eHoltes . 13.Dr. ,Ruinen ,2 k mN W ofth evillag e ofPesse ,Kraloo ;T m17C ,533. 3- 225.3. Area100 0m 2 +abou t50 0m 2. Oak-birch wood with amongst others 105 Trientalis europaea. Coppice,las tcu tabou t1 0year sago ,wit hlow ,mos s covered stubs.Situate d in arelativel y high area ofgroun dmorain ewit h coversand (3L2a), bordering ona broo kvalley .Probabl yplante dbetwee n 1800an d1852 . 14.Dr. , Rolde, 1k mN of the village Schoonloo, Schoonloër strubben; Tm17E , 547.8- 243.1 . Area100 0m 2 +abou t50 0m 2. Oak-birch wood with amongstother s Maianthemum bifolium, Trientalis europaea, and Hedera helix (terrestrial). Coppice,no trecentl y cut,wit hhigh ,mos s coveredstubs . Situated on lowsanddune swit hassociate dplain s (4L8).Th ewoo dwa sfirs t markedo nth e185 2map . 15.Dr. ,Havelte , nearHoltinge r es;T m16H ,534. 9- 213.2 .Are a100 0m 2 + about 1000m 2. Oak-birchwoo d with amongstother s Maianthemum bifolium, Stellaria holostea, and Hedera helix (terrestrial).Coppice ,no trecentl y cut, withhig h stubs.Par to f astri p ofwoodlan dsurroundin gth ecommo n fieldso fth eHoltinge res ,plante di nth e1 9 century. 16.Ov. , Steenwijk, 5k mN ofth evillag e of Steenwijk,Heerlijkhei d De Eese; Tm17E , 537.7- 203.6 . Area 1000m 2 +abou t50 0m2 .Oak-birc hwoo d with amongstother s Maianthemum bifolium and Pteridium aquilinum. Coppice, notrecentl y cut,wit hhig h stubs.Probabl ypar t of astri p ofwoodlan d surroundingth ecommo nfields ,plante di nth e1 9 century. 17. Dr.,Vries .1 k mN W ofth evillag e Zeijen,Zeije r strubben;T m12B , 563.8- 232.1 . Area100 0m 2 +abou t20 0m 2. Oak-brich wood with a great deal of Hedera helix (terrestrial), Maianthemum bifolium, and Pteridium aquilinum. Parto f astri p ofwoodlan dsurroundin gth ecommo nfields ,al readymarke do nth e182 3map . 18.Dr. , Beilen, 1k mS E of the village Hooghalen, Winkelbos; Tm17B , 548.3- 233.3 .Are a70 0m 2. Oak-birch woodwit hamongs tother s Convallaria majalis, Trientalis europaea, and Maianthemum bifolium. Highwood ,lyin go n aridg eo fcoversan d (3L5).Th ewoo doriginate do rwa splante dbetwee n185 2 and1896 ;i ti spossibl ya remnan to fa forme rstri po fwoodlan dsurroundin g thecommo nfields . 19.Dr. ,Gieten ,slightl y N ofth e Zwanemeer;T m12G ,560. 3- 247.2 .Are a 800m 2. High oak-birchwoo d with agrea tdea l of Pteridium aquilinum and Rubus sp.Situate do nth eborde ro fth eHondsru gridge .A nol dwood ,alread y markedo nth e182 3map . 20.Dr. ,Westerbork ,1 k mNN Wo fth evillag eMantinge ,Mantingerbos ;T m17D , 536.5- 236.9 . Area100 0m 2 +abou t50 0m 2. High oak-birch wood, with a great deal of Stellaria holstea and Trientalis europaea. Situated ona raised ground moraine with coversand in abroo kvalley .A nol dwood ,al readymarke do nth e182 3map . 21.Dr. ,Westerbork ,1 k mNN Wo fth evillag eMantinge ,Thijnsbosje ;T m17D , 536.7- 236.7 .Are a50 0m 2.Hig hoak-birc hwoo dwit ha grea tdea lo f Trien talis europaea, Stellaria holostea, and Pteridium aquilinum. Situatedo n groundmorain ewit hcoversand ,borderin ga broo kvalley .A nol dwood ,marke d 106 onth e182 3map . 22.Dr. , Vries, 1k mN W of the village Zeijen, Zeijerstrubben; Tm12B , 563.7- 232.2 . Area87 5m 2 +abou t100 0m 2. High oak-Ilex wood. An old wood,marke do nth e182 3map . 23.Dr. , Norg, Norgerholt; Tm12A , 564.1- 226.3 . Area100 0m 2 +abou t 1000m 2.Hig hoak-Ile xwood .Old ,marke do nth e182 3map . 24.Dr. ,Westerbork ,1 k mNN Wo fth evillag eMantinge ,Mantingerbos ;T m17D , 536.5- 236.9 . Area100 0m 2 +abou t200 0m 2. High oak-Ilex wood. Situated on aris e inth egroun dmorain ei na broo kvalley.'Ol dwood ,marke do nth e 1823map . 25.Dr. ,Beilen , 4k mSS W of the village Wijster,D eHulzedink , Tm17D , 532.9- 232.2 .Are a30 0m 2 (the areabecam e still smallerdurin gthi sin vestigation).Hig h oak-Iiexwood , situated on asmal l rise inth egroun d morainewit hcoversan di na broo kvalley .Ol dwood ,marke do nth e182 3map . 26.Dr. , Hoogeveen, at the Wborde r of thevillag eHoogeveen ,Kinholt s bosje; Tm17C , 527.0- 226.7 . Area100 0m 2. Highoak-Ile xwood , situated ongroun dmorain ewit hcoversand ,borderin ga broo kvalley .Ol dwood ,marke d onth e182 3map . 27.Dr. ,Emmen ,a tth eS Eborde ro fth evillag eEmmen ,alon gth eOevermans - weg;T m17H ,531. 8- 259.1 .Are a80 0m 2.Hig hoak-Iie xwood ,situate do na ridgeo fpossibl ytectoni corigi n (4K1).Ol dwood ,firs tmarke do nth e185 2 map. 28.Dr. ,Westerbork ,1 k mNN Wo fth evillag eMantinge ,Thijnsbosje ;T m17D , 536.7- 236.7 .Are a 500m 2. Highoak-Iie xwood ,situate do ngroun dmorain e with coversand,bordere d by abroo kvalley .Ol dwood ,marke do nth e185 2 map. 29.Dr. , Assen, Asserbos; Tm12D , 556.2- 233.7 . Area of about 500m 2, crisscrosse d or transversedb yman ypath san ddrains .Hig hoak-Ile xwood . Oldwood ,marke do nth e182 3map . 107 ANNOTATIONS ONIDENTIFICATION AND TAXONOMY APPENDIX2 Nomenclature is used according to Moser (1978) for Agericales and Boletales (except for somerecentl ymonographe dgener ath emonograph swer e followed); Donk (1974) for Polyporaceae; Dennis (1978)fo r Ascomycetes; Maas Geesteranus (1975)fo r Hydnaceous fungi;Corne r (1967)fo r Clavarioid fungi;Dumouli n (1968)fo r Gasteromycetes; Eriksson& Ryvarde n (1973-1979) for Corticiaceae;Bourdo t& Galzi n (1927)fo r Heterobasidiomycetes. Many otherbook san darticle swer euse dfo rth eidentifiaction . Material ofman y specieswer econserve d (exsiccata)a tth eherbariu mo f theBiologica l Station,Wijster .Som ecollection sfro mth eRijksherbarium , Leiden,wer estudied ,thes ear eindicate swit h(L) . Colour indications areofte ngive ni ncolou rcodes :Munsel lo rM refer s toth eMunsel lSoi lColou rChart s (1954),Exp ot oth eCod eExpolair e (Cail- leux &Taylo rs.a.) ,an dMethue nt oth eMethue nhandboo ko fcolou r (Kornerup & Wanscher1978) . Amanita porphyria. Det.C .Bas .Spore sspherical . Armillariela mellea. The smallspecie sint owhic hA . meIlea hasbee nspli t areno tdistinguishe dhere . Clitocybe. Many ofth especimen so fth ehygrophanou sspecie so fthi sgenu s weredetermine db yT.W .Kuyper . C. candicans. Often a small,membranaceous , white Clitocybewa s found. Itha d crowded, thingills ,nearl ywithou tsmel lo rtast ean dclearl y belonged to the group ofth e Candicantes. Itgre w onleaves ;spore s weresmall ,ellipsoid , (4.5-)5-6(-6.8 )x 2.9 -3. 9 pm.I twa sdifficul t todecid ewhethe ri twa sC . candicans withlonge rspore so rC . tenuis- sima Romagn.growin go nleaves .I decide dt ofollo wHarmaj a (1969)an d toconside ri ta son especies , C. candicans. Thenam eC . tenuissima is regarded asa younge rsynonym . C. metachroa. Iconside rC . dicolor (Pers. ) Lge and C. metachroa asconspe - cific,an dhav eunite dthe munde rth enam eC . metachroa. C. vibecina. C. langei Sing,e xHor a is included, as Iconside r thema s conspecific. Collybia. Nomenclatureaccordin gt oJanse n& Noordeloo s (1980). Collybiadryophila .Bot hform a funicularis Fr.an df . dryophila werefound . The latterwa sver yvariable :th e colouro fth eca pvarie d fromver y lightt over y darkbrown ,th e formo fth echeilocystidi avarie dfro m more or less cylindrical-coralloid branched, to clavate withthi n branches, to capitate or almost spheropedonculate. They werebot h 108 foundgrowin gterrestria l ando noaktruncs . C. cf kuehneriana. Onlyon especime nwa sfound .I twa sto ool dt odetermin e accurately. Coprinus domesticus, C. micaceus, C. radians andC . xanthothrix areliste d under Coprinus section Micacei intabl e7 an d8 (seeals op.54) . C. tuberosus and C. velox. SeeKit sva nWavere n (1968). Cordyceps spec. 1. Sentt o dr.R.A .Samso n fordetermination ;publicatio n isi npreparation . Cortinarius albofimbriatus Det.P .Ypelaar . C. alboviolaceus. Det.P .Ypelaar . C. bolaris. Det.P .Ypelaar . C. decipiens s.Lge .Det .P .Ypelaar . C. decipiens s.Henry .Ver yofte na rathe rsmall ,brow nTelamoni awit hdis tinct, conicalpapil ,o n arathe r long and slender stemwa s found, which Iconside r as C. decipiens s.Henry .A sther ear ever yfe wpub lications available on this fungus here fellowsa descriptio no fth e specimensstudied . Cap1 5- 3 5m m wide, first convex topulviné , thenplano-conve x to flat, sometimes withundulat e to ascending margin, always with dis tinct, conical papil with rounded top, darkbrown ,dar kredbrown , Munsell2.5Y R2/4-3/6 ,5Y R2/2 ,3/2-4 ,4/3 ,7.5Y R3-4/2-4 ,ofte npale r inmargina l zone to7.5Y R5-6/4 ,hygrophanous ,whe ndryin gturnin gt o browno rpal ebrown ,M 7.5Y R3/2 ,4-6/4 ,10Y R7-8/4 ,whe nyoun gi tha s very fine,whitis hvelu m fibres,whe nolde rthes ear eonl ypresen to n themargi n or iti sglabrous .Gill s (L21-28 ,1 3-7 )slightl yemargi - nate, (3-)4-5(-7)m m high,distant ,abou t8-12/c mhal fwa ymargi nan d cap centre, yellowish ochre,7.5Y R4/4 , 5/6-8, 10YR 5/8, 6/6, edge entire orcrenated ,whe nyoun gwhite rtha nth esides ,whe nol dnearl y the samecolou r asth esides .Ste m (30-)45-75(-90)x2-5mm ,cylindric , base sometimes slightly broader up tox 6mm , occasionaly bent,an d sometimes rooting slightly,wit h fine longitudinal white,silk-lik e shining fibrils,whe nver ywe tcolou r of stem flesh shinesthrough , sometimes it has anindistinc tban d ofwhit evelu m fibreshalfway , footwit hwhite ,occasionall y slightlypin ktomentum ,whic hstick sth e substrate together.Fles h inca p thin,± 1 mm ,i nth epapi lthicker , same colour asth e surface;i n stem thin,slightl y lightertha nth e surface, 5YR4/3 , 5/4,7.5Y R4-5/4 . Smellraphanoi dwhe ncut ,seldo m odourless.Tast esometime sslightl yraphanoid ,usuall yinsignificant . Spores (9.5-)10.2-12.7(-13.9)x(5.4-)6.1-6.8(-8.0) um (approximately 200measurement s in 11collections) , Q1.68-1.76-1.90 , ellipsoid to oblong,ofte nwit h onebi g oildrop,distinctl y rough, the topofte n with some bigger warts, above the apiculus a small,smoothplage . Basidia 30-40x(6-)7-10um , clavate, 4-spored. Pleurocystida absent. Cheilocystidia (15-)20-3lx7.8-10.2(-15.5)um ,clavat et obroa dclavate , 109 notver y distinct fromyoun gbasidia ,wit ha thin ,smooth ,colourles s wall, 1-t omore-celled ,topcel lsometime snearl yspherical ,i nclus ters,i nol dspecimen softe nindistinct ;edg eusuall ypartl ysterile . Habitat.— Inrathe rnutrien tpoo rbu thumu sric hoakwood s onsand y soils,o nan dbetwee nhal frotte nlitter ,clustere dtogether . Collections examined.— Netherlands: prov.Drenthe , Amshoffsbos 10-X-1977,AE J252 ,l-X-1979 ,AE J55 9an d560 ;Tonckensbo s30-X-1978 , AEJ402 ,l-XI-1979 ,AE J587 ;Schoonloë r strubben 19-X-1976,AE J492 , 29-IX-1977, AEJ586 , 22-VIII-1979, AEJ471 ; Schoonloër strubben 3-X-1978,AE J366 ;Heuvingerzan d12-IX-1977 ,AE J220 ;prov .Overijssel , DeEes e 7-IX-1977,AE J491 ,15-IX-1977 ,AE J233 ,26-X-1978 ,AE J400 , 20-VIII-1979,AE J464 ,10-IX-1979 ,AE J529 .(Al lcollection si nHerb . Wag-W). Thebig , large sporesan dth echeilocystidi aar echaracteristic .Bot h characters fit C. decipiens s.Henr ymentione d byKühne r& Romagnes i (1953).Mose r (1978)di dno tmentio n thisfungus .I ti sa grea tpit y thatI wa sno tabl et oconsul tHenry' sorigina ldescription . Inth eWinkelbo s (plot18 )I foun dsom especimen s (8-X-1979,AE J566 ) whichwer ever y similar toth e abovedescribe d c. decipiens butth e papilwa sabsent ,th ecolour swer edarke rred :M 5Y R3/1- 2 whenfresh , and the spores were somewhat smaller: (7.8-)8.3-10.0(-10.8)x(4.9-) 5.4-5.9pm ,Q 1.6-1.9 . Ia mno t sure ifi tbelong st oth esam eo rt o some other species. Intabl e7 i ti sgive nunde r C. decipiens s.Hy , buti tshoul db erea da sc f decipiens. cf fasciatus. InAmshoffsbo s (plot5 ) a funguswa s found(10-X-1977 , AEJ592 )wit hth e following characters:Ca p2 2m m wide,wit hrathe r sharppapi l and ascendingmargin ,hygrophanous ,bleache dt over ypal e brown,Munsel l10Y R8/4 ,papi lbrow n10Y R5/3 .Gill s4 m mhigh ,rathe r distant, yellow, 10YR7/6 . Stem 75x3mm ,hone y coloured, theuppe r half shining and glabrous,th e lower halfwit h somewhit evelu mfi bres.Spore s7.6-8.8(-9.3)x4.9-5. 6 pm (14measurements ,1 collection) , Q1.5-1.6-1.9 , ellipsoid,seldo movoid ,distincl yrough . Refering toLang e (1938)an dKühne r& Romagnes i (1953)i ti seas yt o identifyi ta sC . fasciatus. Itals ofit swit ha descriptio nb yKühne r (1961). The specimenswer erathe rol dan ddrie dwhe nthe ywer efound , soI coul dno tdescrib eth efres hcolours .Thi smake sth edeterminatio n uncertain,s oI hav ecalle dthi sfungu s Cortinarius cf fasciatus. fusispo2~us. Aver yrar efungus ,describe db yKühne ri n195 5an dha ssinc e notbee nmentione d inliterature .Eve nMose r (1978)di dno tkno wthi s species. Ifoun d it5 time s on3 differen tplots ,s oi ti scertainl y not aver y rare species inthi styp eo fvegetation .Description :Ca p 9-29 (-45) mm wide, firstconica l withbroa dinvolute dmargin ,late r becomming half-spherical toconve x to flat,whe n old sometimeswit h ascendingmargin , sometimes with small,fla tumbo ,dar kbrown ,brow n 110 yellow,yello wbrown ,Munsel l 5YR4/6 , 7.5YR4-7/4 ,10Y R6/4 ,finel y radial fibrinöse, first entire later only atmargi n coveredwit h whitish or yellowish velum fibres. Gills up to 5m m high,adnate , reddishbrown , strongbrown ,M 5YR 5/4-6,7.5Y R5/6 , 10YR5/8 ,edg e entiret ofinel ycrenated ,sam ecolou ra sth esides .Ste m17-4 0(-70) x 2-4(-6)mm , more or less cylindric, slightly shining brownyellow , M7.5Y R7/6 ,10Y R6-7/6 ,8/4 ,a t1/ 3 to1/ 2belo wto pwit hsmal lyello w annulus, the lowerhal fwit hsom ewhitis hvelu m fibres,th efoo twit h little,whitis h felt.Fles h inca p andste mwater yyellow ,abou tth e same colour as the surface.Smel l insignificant,whe ncu t sometimes slightlyraphanoid .Spore s (8.0-)8.8-10.7(-12.7)x(4.2-)4'.4-4.9(-5.4) \im (42-measurements in 5coll.), Qmea n 2.2,distinctl y rough,to pwit h bigger warts. Basidia clavate,e.g . 24x7.8pm , 28x8.3jm , 4spored . Gilltramaregular ,hypha ewit hbrow nwall swit hbrow nincruste dpig ment.Habitat. — Innutrient-poor ,humus-poo roakwood so nsand ysoils ; growing on sand.Collection s examined.— Netherlands,prov .Drenthe , Dwingeloo 10-X-1978, AEJ373 ;Zui dHijkerzan d 21-IX-1977,AE J240 , 7-IX-1978,AE J34 2an d343 ;Schoonloë rstrubbe nll-X-1979 ,AE J581 . C. hinnuleus. As Ifoun di tdifficul tt ofin dth erigh tnam efo rthi sspe cies Igiv ea descriptio no fth especimen sstudied .Ca p18-5 6m mwide , firstmor e or lesshalf-spherical , laterbroadl y conical toconvex , with flatumb o andbroad , inflexed margin, darkbrown ,Exp oF- H 52, near marginExp oF 62 ,drie d up yellowishbrown ,Exp oC 63 ,Munsel l 10YR7-8/6 , margin sometimes slightly grayer, indistinctly radially fbirillous,late rnearl ycompletel yfin escaly ,sometime sradia lcleft , marginno tstiate ,wit hsom evelu mfibres .Gill s (L27-32 ,1 3 )5- 9m m high,adnexed ,rathe rdistant ,mor eo rles sth esam ecolou ra sth ecap , yellowishbrown ,M 10Y R 5-6/4.Ste m40-50x5- 9mm ,cylindri co rbroad - eingtoward sth ebas et ox5-l lmm ,a tfirs twhit efibrinou swit habou t halfway adistinc twhite ,persisten tvelu mbel tan dfurthe rdow nman y whitevelu m floes, laterbrowner ,foo twhit e felted,sometime swit ha little lilac. Smell insignificant, slightly nasty-sweetish. Taste slightlyraphanoid-sharp .Spore s (6.3-)7.8-9.0(-9.8)x4.9-5.9(-6.3)Mm , Q1.3 -1. 5 -1.8,ellipsoi dsometime soblong ,distincl yrough ,wit hon e big oildrop.Habitat. — Innutrient -an dhumus-poo roakwood so nsand y soils; on sand. Collections examined.— Netherlands,prov .Drenthe , ZuidHijkerzand ,21-X-1977 ,AE J59 1an dB.W.L .d eVrie s3364 .Th espec imensstudie dcorrespon dt othos edescribe db yHenr y (1936). C. mucifluus Det.P .Ypelaar . C. obtusus. Itwa sals odifficul tt ofin dth erigh tnam efo rthi sspecies . Although the spores are slightly toobi g and adifferen thabita ti s given, these collections agreewit hthos edescribe db yKühne r (1961). Thefollowin gi sa descriptio no fspecimen sfro mth eprovinc eo fOver ijssel. Cap 10-30m m wide, firsthalf-spherica l orbroadl y conical Ill witha rounde dtop ,late rconico-conve xt oconve xo rplano-conve xwit h abroad ,rounde dumbo ,o reve nfla to rwit hslightl yascendin gmargin , darkre dbrown ,Munsel l2.5Y R3/4 ,nea rmargi norang ebrow nt oyellow ish, 5YR 5-6/8,7.5Y R5/6-6/8 ,hygrophanous ,margi nwhe nwe tstriate , withwhit evelu m fibres.Gill s up to 3m m high,a bi tanastomosing , firstligh tbrown ,late rdarker ,yello wbrown ,ochr ebrown ,Munsel l 7.5YR 5-6/8, 5YR5/8 ,toward sedg eslightl yslighter ,7.5Y R7/8 .Ste m 35-45x2-5mm , more or less culindric or slightly attenuatetoward s base, young completelywhit ean dfibrillose ,whe nolde rwhit efibril - losewit h lightbrow n top,late r strongbrown ,7.5Y R5/6 ,wit hsom e white fibres and are dbrow ntop ,2.5Y R4/4 .Fles hi nca pthin ,dar k redbrown , like the surface,i nste m top orangebrown ,5Y R5/8 ,to wardsbas e lighter to7.5Y R6/ 8 infoot .Smel linconspicuous .Spore s (6.6-)7.3-8.3(-9.3)x4.4-5.6 Mm, ellipsoid, sometimes slightly ovoid, indistinctly rough, light yellowish to brownisch yellow underth e microscope. Gilltramayellowis h tobrownis hyello w inKOH ,hymeniu m with some,inconspicuous ,yello wnecropigment .Habitat. — Inoakwood s on nutrient- and humus-poor sandy soils. Collections examined.— Netherland, prov.Overijssel ,D e Eese,2-XII-1976 ,AE J59 4an d595 , 13-X-1978,AE J379 . orellanoides. A beautiful and interesting fungus,tha tI foun donl yi n oneplo ti n2 successiv eyear sa texactl yth esam eplace .Ca p26-3 2m m wide,broadl y conical with arathe r acute top and slightlyinvolut e margin, later convex with distinctly flatumb o and infexedmargin , orangebrown ,yellowis h red,Munse l5Y R5/6 ,7.5.Y R5/6 ,Methue n6D8 , 7E7,no thygrophanous ,completel ycovere db yfin escal yfibril si nth e same colour.Gill s (L± 35 ,1 (0-)l (-3)) up to7 m mhigh ,narrowin g near stem and decurrentb y asmal ltoot h (uncinate),rathe rdistant , ca.12/c mhalfwa ymargi nan dca pcentre ,th esam ecolou ra sth eca po r slightlylighter ,Methue n6D6 ;margi nentire ,th esam ecolou ra ssides . Stem 50-70x6.5-8mm , toward base gradually thickening tox8-1 2m ma t foot,nearl y the same colour asth eca po rslightl ylighter ,Munsel l 5YE5/8 ,7.5Y R6/6-7/8 , abi tlongitudinall y fibrillose,i nlowe rhal f with2 lightyello wvelu mbelts ,10Y R8/6 ,foo tentirel ycovere dwit h lightyello wvelum , 10YR6-8/6 ,2.5 Y 7/6, the tip ofth e foota bi t whitetomentous .Fles hi nca pan dste mligh tyellow ,10Y R8/4 ,turnin g dark brown with KOH. Spores (8.3-)9.3-10.7(-12.7)x(6.3-)6.8-8.3 \im (34measurements ,2 collections) ,Q 1.2 -1.4 -1.6 ,ovoid ,broa dovoid , ellipsoido rbroa dellipsoid ,to psometime sslightl yacute ,distinctl y rough, with one big oildrop.Basidi a (34-)38.5-41.5(-47.3)x8.8-10.7 (-12.2) pm (11measurements) , slenderly clavate,4 spored , old light brown pigmented. Gilltrama consisting of cylindrical to weaklyo r distinctly inflated hyphae, 5-10(-20)|j mwide ,wit h athin ,smooth , light yellowbrow nwal l and some scattered darker spots.Habitat. — 112 In oak-birch woods onnutrient-poo t sandy soils (Querco-Betuletum), between litter and Polijtrichum formosum. Collections examined.— Netherlands,prov .Drenthe ,Amshoffsbo s29-IX-1978 ,AE J359 ,l-X-1979 , AEJ561 . C. paleaceus s.l.Her e Ihav einclude dC . paleaceus Fr.an dC . flexipes Fr. asmentione d by Lange (1938); C. paleaceus Fr.e xWeinm .an dth efor m flexipes Fr.a smentione d by Kühner& Romagnes i (1953);C . paleaceus Fr.an dC . paleiferus Svrceka smentione db yMose r (1978).Lang enote d that C. paleaceus, C. flexipes andC . hemitrichus werever yintimatel y related,speciall y C. paleaceus and C. hemitrichus. C. flexipes should bemor e easilly distinguishable because of the dark gills.Kühne r& Romagnesi dono t refer to Lange'splates ;the y distinguish thevar . flexipes from themor e usual formb yth edar kgills .Mose rconsider s C. paleaceus andC . flexipes s.Lang e synonymous. C. paleiferus is distinguished from C. paleaceus byth eviolaceou sgills .Therefor ei t appears to me to be synonymouswit h C. paleaceus var. flexipes s. Kühner& Romagn . Fora tim eI though tth especimen s Iha dfoun dcoul db edivide dclaer - lyint o C. paleaceus and C. paleiferus. A fungus withviolaeceousl y tomentous stemfoot,violaceou s gills, and aconico-conve x capwit h acutepapill a andman yvelu m floeswa s C. paleiferus.. Amor eslende r funguswit h a flater cap,smalle rpapilla ,mor efugaciou svelum ,an d withoutviolaceou scolour swa s C. paleaceus. Thesiz eo fth espore so f these taxawa s almostth esame .Whil estudyin gth ecollections ,othe r formswer e found:on e similar to C. paleiferus, 2 others similart o C. paleaceus. velum colour of: formo fca p onca p gills topo f tomentumo n stem foot paleiferus conico-convex verymuc h viol viol viol e.g.AE J51 5 bigpapill a persistent cf paleiferus conico-convex fugacious Ibrown viol viol e.g.AE J50 0 smallpapill a cf paleaceus convex fugacious brown brown ±viol e.g.AE J49 6 smallpapill a cf paleaceus convex fugacious brown brown whiteo rnon ( e.g.AE J57 2 smallpapill a footbrow n paleaceus convex much brown brown white e.g.AE J56 2 smallpapill a ±fugacious (scarse) (viol=violaceous;lbrown=ligh tbrown ) Themateria lstudie dwa ss ovarie dtha ti ti spossibl etha tmor ecombi nationsar eprobable .A tthi smomen tI canno tsa ywhethe rthes especi mensbelon gt o1 ,2 ,o rt osevera lspecies .Beside sthis ,i nth efiel d studies "C. paleaceus" wasofte nrecorde dwithou tdetail sbein gnote d aboutcolour s etc.,s o itwa s impossible todistinguis h the5 form s later.I hav etherefo rgroupe dthe mal lunde rth enam eC . paleaceus s.l. 113 C. pluvius. Kühner (1959)describe s C. pluvialis as ane wnam e forC . pluvius s.Orton .Kiihner' sfungu s is,however ,slightl ymor erobus t andha sbigge rspore stha ncoll .AE J521 ;th ecolour sar ecomparitive . Orton'sdescriptio n (1955)i srathe r short,so ,withou tstudyin gth e collections,I d ono tkno wi fthe yar eseparat especies .I hav ecalle d this collection C. pluvius s. Orton. Description: Cap1 1m mwide , broadly conical with arounde dtop ,late ralmos tflat ,crea mcoloure d in centre Methuen 4A2,margi n pure white, surface viscous,a bi t peeling,withou tscales ,fibrils ,o rvelum .Gill s (L± 30 ,1 1-3 )2 m m high, adnate to subdecurrent,rathe r distant,± 12/c mhalfwa ymargi n andca p centre, firstyello w ochre,Methue n4A6 ,late rbrowner ,5C6 . Stem15-20x 2mm , toward base broadering to 3.5mm , subradicating, white,whitish ,a bi tbrownis h4A 3 translucent,solid .Fles h inca p andste mwhite ,i nca pwit hKO Hbrownis hviolaceous .Smel lnast y (Ino- cybe like). Tastever ybitter .Spore s (5.4-)5.9-7.0(-7.3)x4.4-5.4 \m (13measurements) ,ellipsoi do rslightl yovoid ,distinctl yfin erough . Basidia e.g.25x 7pm ,29x6. 8um ,27x6. 3Mm ,slenderl yclavate ,4 spo - red.Cystidi a not seen.Cuticl e consisting ofradia lhyphae ,crowde d together, on septa 5-10M mwide ,inflatin g to 15 MUIwide ,wit hthin , hyalinewalls ,clam p connectionspresen tbu tmostl yindistinct .Habi tat.— In oakwoods onnutrient -an dhumus-poo r sandy soil (Dicrano- Quercetum),betwee nmosses ,especiall y Dicranum scoparium. Collection examined.— Netherlands, prov. Overijssel, DeEese , lO-IX-1979,AE J 521 (2exx.). - C. porphxftopus. Det.P .Ypelaar . C. pseudosalor. Det.P .Ypelaar . C. cf punctatus. (fig.36 )I wa sno t able toge ta clea rpictur e ofC . punctatus fromth eliterature .Especiall ya sth eauthor sdi dno tagre e on the differences between the 3relate d species C. punctatus, C. glandicolor, andC . brunneus. Thecollecte d specimenswer ever ysimila r toC . punctatus s.Lg ean dC . glandicolor s.Rick .Mose r (1978)iden - tifysC . glandicolor asa species fromconiferou swoods .Therefor eI named the specimens C. cf punctatus. Description:Ca p20-35(-50 )m m wide, first conico-convex, later convex with reflexed to slightly ascending, undulatemargin ,wit h abig , acute torounde d umbo,ver y darkbrown ,nearl yblack ,Munsel l 5YR3/1-3 ,10Y R2/1-2 ,margi nwit h radial velum fibres, lighter, grayish brown, 7.5YR5/4 ,10Y R 5/3, hygrophanous, drying toreddis hbrown ,5Y R 5/4 or7.5Y R5-6/4 ,ol d specimens sometimes radial cleft. Gills (L24-32 , 13- 5 (-15))2- 4 (-7) mm high,sometime sveined ,adnat eo rslightl ysubdecurrent ,dis tant,8-10/c mhalfwa ymargi n and centre,rathe rdar kbrown ,greenis h brown, Munsell 5YR4/2-4 , 7.5YR4-5/4 ,5/6 , 10YR4/4-3/3 , 2.5Y4/4 , toward edge sometimes lighter to 10YR6/4 , edge entire orslightl y eroded.Ste m30-50(-70)x(3-)4-6(-9 )mm ,cylindrica lo rslightl yatten - 114 Fig.36 . Cortinarius cf punctatus. Habitsketc h andlongitudina lsections . a and b from A.E.Janse n 475, cfro m A.E.Janse n545 . uateupward ,firs tfinel yfibrinous ,white ,whitish-cream , lightbrown , 2.5Y8/2 ,10Y R8/2 , later darkbrown ,2.5Y R2/4 ,5Y R2/4 ,i no rabov e the middle with indistinct, lightyello wvelu m remains,foo twhit e tomentouswit hviolaceou stint ,no trooting ,soli do rnarro wfistulöse . Smellweakl yraphanoid .Tast einconspicious ,weakl yraphanoi do rweakl y nut-like.Fles h in capver y thinexcep t inumbo , asdar ka sth eca p surface,i nstemcorte x (ca.1 mm )a sdar ka sth eca psurface ,downwar d sometimeslighte rt oyellowbrown ,i nste m"pit ha loose rtissue ,shinin g yellowisht oligh tbrown .Spore s (6.8-)7.8-10.3(-11.2)x(4.4-)4.9-5. 9[im (± 65measurements , 5coll.), ellipsoid orweakl y ovoid, distinctly rough,wit h1 bi goildrop . Basidia (29-)32-35(-39)x6.8-7.8(-8.2) ym, slenderlyclavate ,sometime s constricted under the top,ca .7-1 2 ymprotrudin gabov eth ehymenium , mostly 4-spored with 2.9-3.9|j mlon g sterigmata,ol dbasidi a light yellow brown pigmented. Cheilocystidia inconspicuous, clavate. 115 (13.6-)16-20(-23)x(4.9-)5.9-7.8(-8.3)pm , with thin, hyaline wall, sometimes more-celled or branched and then longer,e.g .26x9. 8 \tm, lying or ascending, alongth esteril eedge .Habitat. — Inoakwood so n nutrient-poor sandy soils (Querco-Betuletum, Violo-Quercetum), in swarms together, under Betula. Collections examined.— Netherlands, prov.Drenthe ,Thijnsbosje ,24-VIII-1979 ,AE J475 ,17-IX-1979 ,AE J539 , 23-IX-1979,AE J545 ;Noordlagerbos ,13-XI-1978 ,AE J405 ;Amshoffsbos , 27-IX-1978,AR J361 . cf stemmatus (fig.37) .Ofte n asmall ,brow n Telamonia wasfound ,tha t was similar to C. stemmatus s.Henry . Henry's fungus,however ,i s somewhatbigger ,ha sbroade r gills,mor evelu m onth e steman dles s variation in the size ofth e spores.Therfor ether e issom edoub t whetherthi swa sreall y C. stemmatus. Description:Ca p (6-)12-22(-35)m m wide, firsthalf-spherical ,conical ,o rbroa d conicalwit hinvolut e margin, later convex,plano-conve x to flato rwit hascendin gmargin , witha big ,rounde do rsometime sacut eumbo ,dar kbrown ,dar kre dbrown , Munsell5Y R 3/4, 4/8, 4/2, 7.5YR 4/2-6, old slightlymor eyellow , 10YR4/3 , 5/4,hygrophanous ,dryin g to7.5Y R5/6 , 6-7/8,10Y R8/4-6 , first entire,late r only atmargi n covered withwhit e orver ypal e brownvelu m fibres,ol dglabrous .Gill s (L± 26 ,1 l-3(-7) )2 m mhigh , adnate to*uncinate, distant,10-12/c mhalfwa y marginan dca pcentre , yellowbrow nt obrown ,7.5Y R4/4 ,5-6/6 ,10Y R5/8 ,edg eth esam ecolou r as the sides.Ste m (10-)20-50x2-4(-6)mm , cylindrical orbroaderin g slightlytoward sth ebase ,t ox6. 5mm ,whitish ,ligh tbrown ,yellowis h brown, Munsell 10YR8-6/4 , 7-6/6, fine longitudinal fibrinous,i n middle with small,whit evelu m ring, lowerdow nsometime ssom escat teredvelu m floes, footslightl yrooting ,wit hlittle ,whit eo rsome times blueish-pink tomentum, which adhers the substrate together. Flesh inca pver ythin ,brown ,darkbrown ,7.5Y R4/2 ,i nste mlighter , 7.5YR4/4 ,10Y R6/6 -8/4 ;wit hKO Hpurplish-black .Smel linconspicuou s orweakl y pelargonium-like atcut .Tast einconspicuous .Spore s (6.6-) 7.3-9.3(-10.2)x(4.4-)4.9-5.6(-6.3)|j m(±8 0measurement s in8 collec tions),Q 1.3-1.6-1.8 ,ellipsoid ,sometime sovoi do roblong ,sometime s withon eo rtw ooildrops .Basidi a26-31x6.3-7. 8 \im, slenderlyclavate , 4 spored,ol dbasidi aligh tt over ydar kbrow npigmented .Cystidi awer e not seen.Gilltram a consistingo fthi ncylindrica lt oinflat ehyphae , 3-15 (jmwide , with brown yellow todar kbrown , finelyt o coarsely incrusted walls. Habitat.— In oakwoods onnutrient - andhumuspoo r sandy soils (Dicrano-Quercetum),i n small tobi gswarm stogether ,o n sandbetwee nmosse s (esp. Dicranum scoparium). Collectionsexamined. — Netherlands, prov.Drenthe , Euidhijkerzand, 21-IX-1977,AE J24 2an d B.W.L.d eVrie s3496 ,7-IX-1978 ,AE J34 4an d345 ,26-I X1978 ,AE J354 , 3-IX-1979,AE J49 5an d501 ,9-X-1979 ,AE J57 1an d573 . 116 Fig.37 . Cortinarius cf stemmatus. Habitsketchs ,an dlongitudina lsection . FromA.E .Janse n495 . C. tabularis. The specimens correspond to.th eplate so fCook e766/78 3an d Lange94B . Cystoderma longisporum. Intabl e7 an d 8 this speciesi scombine dwit hC . amianthinum because theywer eno teas yt odistinguis hmacroscopicall y andwer eno talway sstudie dmicroscopically . Entoloma. Nomenclatureaccordin gt oNoordeloo s (1981). Galerina cf cerina. Foundonl yonce ,th especimen swer eto ool dt ob edeter minedwit hcertainty . G. heimansii. This specieswa s only found once:3 exx .o n arotte ntwig , under Ilex aquifolium in aViolo-Quercetu m ilicetosum,Mantingerbo s (plot24) , 18-X-1978,AE J393 .Th especiemen swer eexactl yth esam ea s thosetha twer edescribe db yReijnder s (1959).Habita tan dfructifica tion-time,however ,wer estikingl ydifferent ! G. cf incurvata. Nom.prov. ,se eBarkma n (1969).Foun donl yonce ,th espe cimenswer eto ool dt ob edetermine dwit hcertainly . G. inversa. Nom.prov. ,se eBarkma n (1969). G.triscopa .(fig .38 )Thi sspecie si sne wt oTh eNetherlands .A descriptio n of thecollectio nAE J27 1 isgiven :Ca p4- 7m mwide ,firs tconical , laterconvex ,wit hdistinc tpapilla ,brownis hyellow ,Munsel l10Y R6/6 , translucentlystiat ewhe nwet ,hygrophanous ,dryin gt oyellow ,2 Y8/6 , cuticleno tpeeling ,margi n slightly longertha ngills .Gill sadnat e tosubdecurrent ,yellowis hbrown ,M 7.5Y R5/8 ,10Y R5/8 ,edg egranular , white. Stem 15-20x0.7-1mm ,yellowis hbrown ,M 7.5Y R5-6/8 ,sometime s reddish yellow near base,M 5Y R5/8 ,th euppe r halfwit h scattered 117 10/u m Fig.38 . Galeriaa triscopa. aSpores ,b Cheilocystidia ,c Caulocystidia ,al l drawings fromA.E .Janse n271 . hairs,th elowe rhal fwit hwhit evelum .Spore s (7.0-)7.8-9.8x4.4-5.4 \im (7measurements) , Q1.5-1.7-1.9 , ovoid, amygdaloid,distinctl yrough , not calyptrate,plag e smooth,to p of spore often smoothwit hsmall , indistinct apical pore, rather dark inKOH ,± 7.5Y R6/6 . Basidia 4 spored.Pleurocystidi aabsent .Cheilocysidi a32-44x5.4-8. 3 (ventricose base)x(2.0-)2.9-3.9(-4.6) (neck)x(2.9-)3.4-6.1(apex )pm ,lageniform , ventricose, with rather long neck, narrowing towards the top,to p usually with distinct spherical capitulum, sometimes onlyrounded ; abundant,edg e sterile.Caulocystidi a slightlysmaller ,approximatel y thesam efor ma sth echeilocystidia .Habitat. — Onrotte nwoo di noak - 118 birchwood onnutrient-poo r sandysoil s (Querco-Betuletum).Collectio n examined.— Netherlands, Prov.Drenthe , Noordlagerbos, 25-X-1977, AEJ271, .Foun d at the same locality on 27-IX-1976, AEJ162 ,an d ll-X-1978,AE J377 . When these specimens were compared to the literature (e.g.Kühner , 1935,Smit h& Singer ,1964 )an dothe rcollection so f G. triscopa (BRD, Eifel, Gerolstein, 4-X-1971, C.Bas571 8 (L)the y were smaller and somewhat lighter, more yellowish, coloured. Microscopical features wereth esam ea sthos ei nliteratur ean di nth ecollectio nC B5718 . Hebei omac f vaccinum. Foundonl yonce :o nplo t15 ,Dr. ,Uffelte ,27-IX-1979 , AEJ553 ,2 exx .Th especimen swer esimila rt o H. vaccinum, butdiffere d slightly: the capwa s umbonate,th e stemwa sbulbou s (in1 specime n only), the gill-edge beared some drops on it,an d thebasidi a and cheilocystidiawer e slightly shorter.Nevertheless , H. vaccinum seems tob eth ecorrec tname . Inocybelongicystis . Cheilo- and pleurocystidia broadly fusiform, 65-73x 15-22Mm . I. ovatocystis. Cheilo-an dpleurocystidi a21-43x13-2 1|jm . I. sambucina. The specimenscorresponde dwit hth edescriptio nan dplat eb y Bruylants (1957). Inocybe xanthomelas. fig.39 .Description :Ca p14-3 0m mwide ,firs tconical , later broadly conical to convex or flat with slightly undulating margin, with distinctumbo ,whe nol d sometimes with aradia l cleft margin,ligh tyello wbrown ,yello wbrow nt obrown ,Munsel l2.5Y R7/6 , 10YR6/4-6 , 7.5YR6/6 , 5/4-8,4/4 , rathercoars e radial fibrinöse. Gills ±2. 5m m high, adnexed, rather crowded,whitis h to lightyel lowishgray ,M 2.5 Y7/2 ,late rbrowner ,10Y R6-7/4-6 . Stem15-40x2.5- 4 mm,cylindrica lo rslightl ybroaderin gtoward sth ebase ,th efoo twit h asmal lt orathe rrobust ,marginat ebulb ,firs tnearl ywhite ,becomin g light yellow, M5 Y8/4 , 2.5Y8/2-6 , later more brown, 2.5Y7/4-6 , 10YR5/3 , 6/3-4, fine fibrinöse, thewhol e lengthpruinose ;solid . Flesh inca pwhite ,i n stemver y lightyellow , inbul bwhite .Smel l and taste inconspicuous. Spores (7.8-)8.8-10.7(-11.2)x(5.9-)6.3-7.8 (-8.3)p m (41measurements , 5collections) , Q1.2-1.4-1.6 , with (7-)8-12(-15)prominen t noduls. Basidia 26-36x8.2-12.2 \im, clavate, 4 sporedwit h up to 6p m longsterigmata .Pleurocystidi a 51-92x11-20 (-25) \im, lageniform, seldom clavate to spheropedonculate, with a 2.5-3.4(-4.4)p mthick ,i nKO Hyellow ,smoot hwall ,th eto pofte nwit h somecrystals .Cheilocystidi a from about same form and size.Caulo - cystidia lageniform, (52-)70-100(-112)xl2-24Mm ,wit ha thick ,i nKO H yellow wall,~eve n thicker atth eneck ,betwee nles sstrikin gclavat e cells.Habitat.- -I noakwood s onnutrien tan dhumu spoo rsand ysoils , in small swarms on bare sand or between mosses such as Dicranum scoparium, Pohlia nutans. Collections examined.— Netherlands,prov . 119 <^r^> Fig.39 . Inocybe xanthome las. a Habit sketchs and longitudinal section, bSpores , cPleuro -an d cheilocystidia. a fromA.E .Janse n499 , ban dc fro mA.E .Janse n189 . 120 Drenthe, ZuidHijkerzand , 28-X-1976,AE J284 ,19-VI11-1977 ,AE J189 , 26-VII-1978,AE J304 ,3-IX-1979 , AEJ499 ,9-X-1979 , AEJ 570;Schoon - loërstrubbe nll-X-1979 ,AE J579 ;prov .Overijssel ,D eEese ,10-X-1979 , AEJ513 .Observations. —Th e specimens studied were largely fromon e plot,wher e theywer e found ineac ho f the4 years ,a texactl yth e same location;ther ewer eprobabl y 3myceli a intha t 1000m l sample plotarea . Therewa s somevariatio ni nmicroscopi c features.Collection s30 4an d 570 had comparatively small spores, 7.8-9.3x6.3-6.8 \im resp. 8.8-9.3x5.9-6.8(-7.8)|jm , whereas 189 has 9.3-11.2x6.2-8.3 Mm. Collection 570ha dbot h short and thick, and long and thinpleuro - cystidia (Q2.0-7.3 ,mea n4.0 )wherea sothe rcollection shav elon gan d thinpleurocystidi a with lessvariatio n (e.g.Q 3.1-4.8 ,mea n3. 9o r 4.1-5.2,mea n4.6) . Distinguishing /. xanthomelas and I. mixtilis (Britz.)Sacc .wa sno t easy. I. xanthomelas shouldhav ea blackenin gstem ,bigge rspore swit h more prominent noduls and more slenderpleuro -an dcheilocystidia , thanJ . mixtilis. Inth ecollection s studied Ineve robserve dblack ening ofth e stemwhe ndrying ,bu t aged specimens inth e fieldha d brownishstems .Spore swer emostl yo fth efor man dsiz eappropriat et o I. xanthomelas, buti nsom ecollection sth esiz ewa smor elik etha to f I. mixtilis. Form and sizeo fth epleuro -an dcheilocystidi awas ,i n spite of the variation, more liketha to f I. xanthomelas, butth e thickening of thewal lwa smor e liketha to f I. mixtilis (seeStang l 1977 and 1980). Iti smos tlikel ytha tth especimen sstudie dbelonge d to I. xanthomelas, butther ewer esom edifferences . Marasmiellus. Nomenclatureaccordin gt oJanse n& Noordeloo s (1980). Psathyrella. Many of the specimens were determined by dr.E .Kit sva n Waveren. P. fulvescens var. dicrani. Fig.40 :A smal l Psathyrella wasofte nfoun di n theDicrano-Quercetum . Itha d ahygrophanous ,war m orangebrow nca p with fugacious,whit evelu man da slender ,no nrootin gstem .A tfirs t I thought itwa sa for mo fP . prona. Butaccordin gt odr .E .Kit sva n Waveren,wh owa s sokin d ast o study someo fth especimens ,i ti sa formo f P. fulvescens. P.' fulvescens isa ver yvariabl especie san di t isa tpresen tdifficul tt odescrib ene wvarietie so rforms .Neverthe less itseem s tom etha tthi sfor mha ssom econstan tdifference swit h thetypica l form. Itha sbigge r spores,strikin gcheilocystidi awit h often bent tops, andha s onlybee n found inth eDicrano-Quercetum , never inth emor ehumus-ric h oakwoods.Therefo r Iwan tt ogiv ei ta taxonomicalran kan da si ti sno tye tclea rwhethe rintermediate soccu r ornot ,I giv ei tth elo wran ko fvariety . Psathyrella fulvescens var. dicrani A.E.Jansen ,var .nov .Fig .40 .A varietate typica differt sporibusmajoribus ,(8.3-)9.3-11.4(-12.2 )p m 121 longis, (4.9-)5.5-6.1(-6.8) \im latis, cheilocystibus collo paulum gracilioreapicegu eseap ecurvato . Typus. The Netherlands, Prov. ofDrenthe ,Zui dHijkerzand ,2 6jul e 1978,A.E .Janse n30 3 (Herb.Wag-W) . Description: Cap (10-)18-30(-40)m m wide,conico-conve x toconve xwit ha saml lt o rather stout,rounde d umbo,rathe r dark redbrow ni ncentre ,Munsel l 5YR4/3-8 , 7.5YR4/4 ,lighte r nearth emargin ,5Y R5/3 ,7.5Y R5/4-6 , hygrophanous,guickl ybecomin g lighter incolour ,war morang ebrown , 7.5YR5-7/6 ,5/8 ,10Y R7-8/ 6 incentre ,nea rmargi nmor elight ,brown ish-yellow,ofte nwit h somepink ,10Y R7/3 , 6-8/4, 6-7/6,6/8 , 2.5Y 8/4, firstwit h distinctwhite ,fugaciou svelu m atth emargi nsome timeswit h fine floescoverin ghal fo fth ecap ,whe nolde rwit honl y some floesnea rmargi n or glabrous.Gill s (L20-35 , 11-3 ) 3-5.5m m broad,adnate ,ver ythi nan dtherefor erathe rdistant ,ca .16/c mhalf waymargi nan dca pcentre ,firs tyello wbrown ,Munsel l10Y R7/6 ,late r with spores grayish brown,lightgray ,brow ngra y todar kchocolat e brown,2.5Y R5-6/2 ,10Y R6/1-3 ,4/2 ,3/3 ,edg ewhit ebu tnea rca pmar ginofte nth e samecolou ra sth esides .Ste m30-60x2-4(-5 )mm ,cylin drical,sometime snea rbasi sbecomin gslightl ybroade rt ox3-7(-9 )mm , white,seldo myello wwhit e orligh tbrownis hwhite ,to ppruinous ,i n lower half fine fribrillous and with some velum fibres,whe nol d glabrous, foot rounded, notrooting , fistulöse.Fles h inca pthin , darkyello wbrow n to lightbrow n orcream ,10Y R4-8/4 ;i nste mwhit e or creamy white at thetop ,ligh tyello wbrow n 10YR8/ 6 roundth e fistula,yello wbrow n 10YR5/ 6 atth e foot;i ngil lhyalin et oligh t yellow brown. Smell and taste inconspicious.Spore s (8.3-)9.3-11.4 (-12.2)x(4.9-)5.5-6.1(-6.8)|j m (±5 0measurements , 9 collections), Q1.4-1.7-1.9 , ellipsoid, with small pore, dark browngra y inKO H under microscope..Basidi a ca.20-27xi oMm , broad clavate to almost spheropedunculate, 4spored . Pleurocystidia 36-60x10-14 (ventricose base)M"> . fusiform-ventricose or lageniform-pedicellate with long, tapering neckwit h subacutetop ,wit h aligh tyello w (inKOH) ,thi n wall. Chailocystidia ofalmos tth esam efor man dsize ,bu tnec kofte n more slender andmor edistinctl y separated fromventricos ebase ,to p oftenbent ,mixe dwit hfusifor m andspheropedunculat e cells,formin ga sterile edge.Habitat. — Solitary ori nsmal lgroups ,o nlitte ro ro n sandbetwee nmosse s sucha s Dicranum scoparium, Pohlia nutans, Campu- lopus flexuosus, in oakwoods onnutrien t andhumu spoo r sandysoil s (Dicrano-Quercetum). Collections examined.— Netherlands, prov. Drenthe, ZuidHijkerzand , 9-XI-1972P .Ypelaa r72 ,28-X-197 6AE J455 , ll-V-1977 AEJ170 , 19-VIII-1977 AEJ456 , 21-IX-1977 B.W.L.d eVrie s 109-1, 26-VII-1978 AEJ303 , 9-VIII-1978AE J316 , 7-IX-1978AE J346 , 348an d349 ,3-IX-197 9 AEJ494 ,9-X-197 9AE J569 ;Boswachteri jDwinge - 122 *> '--.- 40 um • • • Fig. 40. Psathyrella fulvescezis var. dicrani. aan db Habi tSketch s (x0.7)' andlongitudina lsection s (xo.4),c an dd Spores ,e Pleurocystidia , fan dg Cheilocystidia .a fro mA.E .Janse n494 ,b fromA.E .Janse n 569, can df fro mA.E .Janse n170 ,d ,e ,an dg fro mA.E .Janse n303 . 123 loo, 25-VII-1960 A.K.Masselin k 6543, 19-VIII-1972 P.Ypelaa r4 , 8-V-1977AE J168 ,10-X-197 8AE J372 ;Boswachteri jSmilde ,10-VIII-197 3 P.Ypelaa r96 ;prov .Overijssel ,D eEese ,2-XII-1976 ,AE J157 . Russula emetica. Thevarietie s sylvestris Sing., betularum (Hora)Romagn . (=R. betularum Hora), and emetica were found. Ihav eno tstudie dal l specimens microscopically, which isnecessar y inorde rt onam eth e varietiescorrectly ,s oI conside rthe mher ea s R. emetica s.l. Stropharia aeruginosa was considered including S. caerulea Kreisel(=S . cyanea (Bolt.ex.Fr. )Tuomikoski) . Steccherinum cf hydneum. Specimens differed from S. laeticolor (Berk.& Curt.)Bank . (-S. robustius Eriks.& Lundell )i nth emor eochraceou s carpophore and in thenearl y subglobosespores :4-4.6(-5.0)x3- 4 \m. Det.N .Deodatu san dB.W.L .d eVries . 124 INDEX TO THE FUNGI (Glmean stabl e7 an d8 grou p 1,G 2mean stabl e7 an d8 grou p2 ,etc. ,onl y indicatedi nfung ino tmentione di nth etext. ) Amanita citrina (Schaeff.)S.F .Gra y57 ,5 9 A. fulva (Schaeff.ex )Pers .56 ,8 1 A. muscaria (L.e xFr. )Hooke rG l A. porphyria (A.& S .e xFr. )Seer .53 ,10 7 A. rubescens (Pers.e xFr. )S.F .Gra y7 7 Armillariella mellea (Vahl.e xFr. )Karst ,s.l .63 ,73 ,77 ,90 ,93 ,97 ,10 7 Bjerkandera adusta (Wildpe rFr. )Karst .54 ,5 8 Boletus edulis Bull,e xFr .5 6 B. erythropus (Fr.ex .Fr. )Pers .5 7 Bulgaria inquinans Fr.G 2 Calocera cornea Batsch77 ,8 3 C. viscosa Pers.G 2 Cantharellus cibarius Fr.5 7 C. cibarius var. amethysteus Quel.5 3 C. sinuosus (Fr.)Kühn .& Romagn .5 3 Chondrostereum purpureum (Pers.e xFr. )Pouz .5 8 Ciboria batschiana (Zopf)Buchw .49 ,6 1 Clavariadelphus junceus (A.& S .e xFr. )Corne r5 4 Clavulina cristata (Fr.)Schroet .G 2 Clitocybe angustissima (Lasch.)Kumm .5 3 C. brumalis (Fr.e xFr. )Kumm .5 9 C. candicans (Pers.e xFr. )Kumm .62 ,10 7 C. cerrusata (Fr.)Kumm .5 3 C. clavipes (Pers.e xFr. )Kumm .G 7 C. diatreta (Fr.e xFr. )Kumm .5 8 C. ditopa (Fr.e xFr. )Gill .G 7 C. flaccida (Sow.e xFr. )Kumm .60 ,61 ,77 ,7 8 C. fragrans (Sow.e xFr. )Kumm .5 8 Clitocybe gibba (Pers.e xFr. )Kumm .= C . infundibuliformis (Schaeff.ex .Fr. )Quel .5 3 C. inornata (Sow.e xFr. )Gill .G l C. lohjaënsis Harm.5 3 C. metachroa (Fr.)Kumm .58 ,10 7 C. odora (Bull,e x Fr.) Kumm .5 3 C. phyllophila (Fr.)Quel .G 2 125 C. umbilicata (Schaeff.e xFr. )Sing .G 2 C. vibecina (Fr.)Quél .63 ,69 ,83 ,10 7 Clitopilus hobsonii (Berk.& Br. )Orto n5 8 Collybia butyracea (Buil.e xFr. )Kumm .62 ,77 ,81 ,8 7 C. cirrhata (Schuin,e xFr. )Kumm .62 ,7 7 C. cookei (Bres.)J.D .Arnol d58 ,90 ,93 ,9 7 C. dryophila (Buil.e xFr. )Kumm .10 7 C. fusipes (Buil.e xFr. )Quél .59 ,60 ,9 2 C.cf . kuehneriana Sing.= Marasmius bresadolae s.Kühn .& Romagn .53 ,10 7 C. maculât a(A .& S .e xFr. )Kumm .G 8 C. peronata (Bolt.e xFr. )Kumm .60 ,77 ,79 ,83 ,8 7 C. tuberosa (Bull,e xFr. )Kumm .9 3 Coltricia perennis (L.pe rFr. )Murr .5 3 Coprinus heterosetulosus Locg.5 3 C. domesticus (Boit,e xFr. )S.F .Gra ys .Métrod. ,Kühn .& Romagn .54 ,6 1 C. micaceus (Bull,e xFr. )Fr .54 ,6 1 C. pellucidus Karst.G 2 C. radians Desm.54 ,6 1 C. stellatus Bullerapu dBisb y5 3 C. tuberosus Quél.= C. stercorarius (Buller)Fr .s .Lg e10 8 C. velox Godeyapu dGille t10 8 C. xanthothrix Romagn.54 ,6 1 Coriolus versicolor (L.pe rFr. )Quél .G 9 Cordyceps canadensis Ellis& Everhar t5 7 C. militaris (L.e xSt.-Am. )Lin k5 3 C. ophioglossoides (Ehrenb.e xFr. )Lin k57 ,8 1 C.spec . 153 ,10 8 Cortinarius albofimbriatus Henry10 8 C. alboviolaceus (Pers.e xFr. )Fr .57 ,10 8 C. bolaris (Pers.e xFr. )Fr .57 ,10 8 Cortinarius decipiens s.Lge .10 8 C. decipiens s.Henr y108 ,10 9 C. delibutus Fr.5 3 C. elatior Fr.G 7 C.c f fasciatus (Scop.)Fr .53 ,10 9 C. fusisporus Kühn.57 ,10 9 C. glandicolor (Fr.)Fr .5 7 C. henitrichus (Pers.e xFr. )Fr .5 3 C. binnuleus Sow.e xFr .53 ,11 0 C. mucifluus s.Rick. ,Konr .& Maubl. ,no nFr .11 0 C. obtusus (Fr.)s .Henr y57 ,110 ,11 1 C. orellanoides Henry59 ,11 1 C. paleaceus Fr.s.l .57 ,11 2 C. pluvius (Fr.)Fr .s .Orto n53 ,11 3 126 C. porphyropus (A.& S. )Fr .11 3 C. pseudosalor J.Lang e53 ,11 3 C.c f punctatus s.Lang e113 ,11 4 C.c f stemmatus Fr.s .Henr y57 ,115 ,11 6 C. tabularis (Bull,e xFr. )Fr .61 ,11 6 Crepidotus haustellaris (Fr.e xFr. )Kumm .9 2 C. pubescens Bres.58 ,83 ,9 3 C. variabilis (Pers.e xFr. )Kumm .G 8 Crucibulum laeve 53 Coryne sarcoides (Jacq.e xS.F .Gray )Tulasn eG 9 Cudoniella acicularis (Bull,e xFr. )Schroet .49 ,58 ,7 7 Cyathipodia macropus (Pers.e xFr. )Denni s5 3 Cystoderma amianthinum (Scop.e xFr. )Fay .54 ,62 ,11 6 C. longisporvm (Kühn.)Heinem .& Thoe n54 ,11 6 C. granulosum (Batsche xFr. )Fay .5 3 Daedaleopsis confragosa (Bolt,pe rFr. )Schroet .5 3 Dermocybe cinnamomeolutea (Orton)Mos .5 7 D. crocea (Schaeff.e xFr. )Mos .5 3 Entöl oma cetratum (Fr.)Mos .G l E. euchrous (Pers.e xFr. )Don k60 ,9 2 E. farinogustus Arnolds& Noordeloo s5 3 E. rhodocylix (Lasch)Mos .G 7 E. turbidum (Fr.)Quel .5 7 Exidia glandulosa (Bull.)Fr .4 9 Fistulina hepatica (Schaeff.)pe rFr .G 8 Flammulaster carpophiloides (Kühn.)Watl .5 3 Fomes fomentarius (L.e xFr. )Fr .5 3 Galerina allospora Sm.& Sing .= G . luteofulva OrtonG 9 C. ampullaceocystis Orton59 ,60 ,9 2 G. atkinsoniana Smith62 ,8 1 G. calyptrata Orton57 ,59 ,77 ,8 1 G.c f cerina Sm.& Sing .53 ,11 6 G. cinctula Orton58 ,9 2 G. decipiens Sm.& Sing .81 ,9 6 G. heimansii Reijnd.54 ,11 6 G. heterocystis (Atk.)Sm .& Sing .5 9 G. hypnorum(Schran ke xFr. )Kühn .5 8 G.c f incurvata Barkman53 ,11 6 G. inversa Barkman53 ,11 6 G. mniophila (Lasch.)Kühn .G 7 G. pumila (Pers.e xFr. )M .Lg ee xSing .G 8 G. sahleri (Quel.)Kühn .5 8 G. triscopa (Fr.Kühn .59 ,92 ,116 ,117 ,11 8 Ganodermaapplanation (pers.pe rS.F .Gray )Pat .G 9 127 Gymnopilus hybridus (Fr.e xFr. )Sing .77 ,8 1 G. spectabilis (Fr.)Sing .G 2 Hapalopilus rutilans (Pers.pe rFr. )Karst .= H. nidulans (Fr.) Karst. 58,77 , 80 Hebeloma calyptrosporum BruchetG 7 H. crustuliniforme (Bull,e xSt.-Am. )Quel .5 3 H. longicaudum (pers.e xFr. )Kumm .G 7 H. pumilum J.Lg e5 7 H.c f vaccinum Romagn.53 ,11 8 H. velutipes Bruchet5 3 Selvella crispa Fr.5 3 Heterobasidion annosum (Fr.)Bref .= Fomes annosus (Fr.)Karst .9 3 Hohenbuehelia atrocaerulea (Fr.e xFr. )Sing .5 8 Hydnellum compactum(Pers .ex .Fr. )Karst .5 3 H. concrescens (Pers.e xSchw. )Banke r= H. velutinum var. zonatum (Fr.) MaasG .G l H. scrobiculatum (Fr.e xSecr. )Karst .5 7 H. spongiosipes (Peclc)Pouz .= H. velutinum var. spongiosipes (Peck) MaasG .5 7 Hymenoscyphus epiphyllus (Pers.e xFr. )Rehm .ap .Kau ffin. 49 ,5 9 H. fructigenus (Bull,e xMérat )S.F .Gra y4 9 Hypholoma fasciculare (Huds.e xFr. )Kumm .73 ,77 ,9 0 H. sublateritium (Fr.)Quel .58 ,7 7 Incrustoporia semipileata (Peck)Don k5 3 Inocybe boltonii HeimG l I. cookei Bres.G 9 I. lacera (Fr.)Kumm .5 7 I. lanuginella (Schroet.ap .Cohn )Konr .& Maubl .5 3 I. longicystis Atk.= I. lanuginosa (Bull,e xFr. )Kumm .var . lanuginosa stangl11 8 I. margaritispora (Berk.ap .Cke. )Sacc .5 3 I. napipes J.Lg e57 ,77 ,81 ,9 6 I. ovatocystis Kühn.57 ,81 ,11 8 I. petiginosa (Fr.e xFr. )Gill .5 3 I. sambucina (Fr.)Quel .57 ,11 8 I. xanthomelas Bours.&Kühn .= I. xanthomelaena s. Kühn.& Romagn . 57,81 ,118 ,119 ,12 0 Inonotus dryadeus (Pers.e xFr. )Murr .6 0 I. polymorphus (Rostk.)Pilâ ts .Bourd. ,L .Mair e5 3 I. radiatus (Sow.pe rFr. )Karst .5 8 Kuehneromyces mutabilis (Schaeff.e xFr. )Sing .& Sm .5 8 Laccaria amethystina (Bolt,e xHook. )Murr .57 ,59 ,62 ,7 7 L. laccata (Scop,e xFr. )Bk .& Br .5 8 I. proxima (Boud.)Pat .6 2 128 lactarius camphoratus (Bull,e xFr. )Fr .57 ,59 ,7 7 L. chrysorheus Fr.57 ,77 ,78 ,8 1 L. quietus (Fr.)Fr .63 ,77 ,8 7 h. theiogalus (Bull,e xFr. )S.F .Gra y= L. tabidus Fr.63 ,7 7 L. turpis (Weinm.)Fr .= L. necator (Bull.em .Pers .e xFr. )Karst .G 8 Laetiporus sulphureus (Bull,pe rFr. )Murr .6 0 Lentinellus cochleatus (Pers.e xFr. )Karst .G 7 Leotia lubrica Pers.5 7 Lepiota cristata (A.& S .e xFr. )Kumm .5 3 Lepista gilva (Pers.e xFr. )Roz eG 7 L. nuda (Bull,e xFr. )Cke .9 2 Leucocoprinus brebissonii (Godeyap .Gill. )Loq .5 4 Lucoperdon foetidum Bonord.G 8 L. spadiceum Pers.5 3 Macrotyphula fistulosa (Holmskj.e xFr. )Peterse n5 4 Marasmiellus ramealis (Bull,e xFr. )Sing .58 ,8 3 ». vaillantii (Pers.e xFr. )Sing .= » . languidus (Lasch)Sing .12 0 Marasmius androsaceus (L.e xFr. )Fr .57 ,77 ,81 ,9 6 ». epiphylloides (Rea)Sacc .& Trott .60 ,61 ,77 ,78 ,83 ,87 ,9 6 ». splachnoides (Fr.)Fr .6 0 Melanoleuca melaleuca (Pers.e xFr. )Murr .5 3 Merulius tremellosus Fr.49 ,5 8 Mutinus caninus (Huds.e xPers. )Fr .6 1 Mycena capillaris (Schum.e xFr. )Kumm .G 2 ». cinerella Karst.69 ,77 ,83 ,8 7 ». epipterygia (Scop.e xFr. )S.F .Gra y5 8 ». galericulata (Scop.e xFr. )S.F .Gra y9 2 ». galopus (Pers.e xFr. )Kumm .8 1 ». haematopus (Pers.e xFr. )Kumm .58 ,9 2 ». hemisphaerica Peck5 4 ». inclinata (Fr.)Quel .59 ,60 ,9 2 ». iodiolens Lund.s .Kühn .& Romagn . (=» . amygdalina (Pers.)Sing. ? accordingt oMose r1978 )G 9 ». mucor (Batsche xFr. )Gill .5 8 ». oortiana Kühn,e xHor a5 4 ». peasoniana Dennise xSing .= » . pseudopura s.Kühn ,no nCke .6 0 ». polyadelpha (Lasch)Kühn .5 3 ». polygrama (Bull,e xFr. )S.F .Gra y9 2 ». pura (Pers.ex .Fr. )Kumm .5 8 ». rorida (Scop.e xFr. )Quel .59 ,77 ,9 3 M. sanguinolenta (A.& S .e xFr. )Kumm .63 ,7 7 ». sepia J.Lge .= » . vitrea s.Kühn . &••Romagn .5 8 ». smithiana Kühn.G 2 ». speirea (Fr.e xFr. )Gill .G 9 129 Ä. stylobates (Pers.e xFr. )Kumm .58 ,7 7 M. vitilis (Fr.)Que l= M. fHopes s.Kühn .& Romagn .58 ,8 7 Nectria sp.7 7 Onygena corviaa A.& S .e xFr .49 ,9 3 Oudemansiella platyphylla (Pers.e xFr. )Mos .58 ,69 ,77 ,8 3 Panaeolus fimicola (Fr.)Gill .5 3 Panellus serotinus (Pers.e xFr. )Kühn .58 ,8 7 P. stipticus (Bull,e xFr. )Karst .5 9 Paxillus involutus (Batsch.)Fr .63 ,77 ,81 ,87 ,90 ,9 1 Phallus impudicus L.e xPers .58 ,73 ,77 ,82 ,9 1 Phellinus ferreus (Pers.)Bourd .& G .54 ,6 1 Phellodon confluens (pers.)Pouz .= Hydnum confluens Pers.5 3 P. melaleucus (Sw.ap .Fr .e xFr. )Karst .G l P. niger (Fr.e xFr. )Karst .5 3 Phlebia radiata Fr.49 ,8 3 P. rufa (Fr.)Christ .49 ,5 4 Pholiota alnicola (Fr.)Sing .9 2 P. lenta (Pers. ex Fr. ) Sing. G8 P. tuberculosa (Schaeff.e xFr. )Kumm .9 2 Phylloporus rhodoxanthus (Schw.)Bres .5 4 Piptoporus betulinus (Bull,pe rFr. )Karst .5 8 Pleurotus dryinus (Pers.ex .Fr. )Kumm .5 4 P. ostreatus (Jacg.ex .Fr. )Kumm .5 8 Pluteus cervinus (Schaeff.e xFr. )Kumm .73 ,83 ,8 7 P. salicinus (Pers.e xFr. )Kumm .5 8 Polyporvs brumalis (Pers.)pe rFr .58 ,77 ,8 0 P. ciliatus (Fr.)pe rFr .5 4 P. varius (Pers.)pe rFr .6 1 Psathyrella cernua (Vahle xFr. )ap .Gam sG l P. cotonea (Quel.)Konr .& Maubl .5 3 P. frustulenta (Fr.)A.H .Sm .6 1 P. fulvescens (Romagn.)Mos .8 1 P. fulvescens var. dicrani A.E.Janse n57 ,58 ,97 ,120 ,12 1 P. fusca (Schum.e xFr. )Mos .ap .Garn s5 3 P. gossypina (Bull,e xFr. )Pearso n& Denni s5 3 P. hydrophila (Bull,e xMérat )Mair e58 ,9 2 P. cf obtusata (Fr.)A.H.Sm .5 3 P. spadiceogrisea (Fr.)Mair eG 9 P. squamosa (Karst.)Mos .ap .Gam s7 7 Psilocybe crobula (Fr.)M .Lg ee xSing .58 ,77 ,8 9 P. inguilina (Fr.e xFr. )Bres .5 3 Ramaria flaccida (Fr.)Rick .5 3 Resupinatus applicatus (Batche xFr. )S.F .Gra y6 0 Rickenella fibula (Bull,e xFr. )Raith .= Mycena fibula (Bull,e xFr. ) Kühn.5 8 130 fi. setipes (Fr.)Raith .= Mycena swartzii (Fr.e xFr. )A.H.Sm .5 3 Ripartites helomorpha (Fr. ) Karst. 53 Russuia adusta (Pers.e xFr. )Fr .5 7 R. aeruginea Lindbl. ex Fr. 53 R. amoenolens Romagn.G 9 R. atropurpurea (Krombh.)Britz .G 8 R. brunneoviolacea Crawsh.5 3 R. cyanoxantha (Schaeff.e xSecr. )Fr .5 7 R. cuanoxantha f. peltereaui Maire5 3 R. emetica (Schaeff.e xFr. )S.F .Gra y62 ,81 ,12 3 fi. fellea Fr.5 4 R. fragilis (Pers.e xFr. )Fr .57 ,8 1 Ä. heterophylla (Fr.)Fr .5 3 fi. laurocerasi Melzer5 3 R. mairei Sing.5 3 fi. nigricans (Bull,e xMérat )Fr .G 9 fi. nitida (Pers.e xFr. )Fr .5 3 fi. ochroleuca (Pers.e xSecr. )Fr .G 9 fi. parazurea J.Schaeff .6 0 fi. raoultii Quel.5 3 fi. vesca Fr.5 7 Rutstroemia firma (Pers.)Karst .8 3 Sarcodon joeides (Pass.)Bat .G l S. scabrosus (Fr.)Karst .5 7 S. underwoodii Banker53 ,5 7 Schizopora paradoxa (Schrad.pe rFr. )Don kG 9 Scleroderma areolatum Ehrenb.5 3 S. citrinum Pers.77 ,7 9 S. verrucosum Bull,tran sPers .60 ,6 1 Sphaerobolus stellatus Todepe rPers .4 9 Steccherinum cf hydneum (Rick.)Maa sG .12 3 Stereum gausapatum 49, 54 S. hirsutum (Wild,e xFr. )S.F .Gra y49 ,5 8 S. rugosum(Pers .e xFr. )Fr .49 ,58 ,8 3 Stropharia aeruginosa (Curt,e xFr. )Quel .59 ,12 3 S. semiglobata (Batsche xFr. )Quel .G 8 Telephora terrestris Fr.57 ,8 1 Tephrocybe ambusta (Fr.e xFr. )Don kG 9 T. tylicolor (Fr.e xFr. )Mos .= T. tesquorum (Fr.) Mos. = Lyophyllum plexipes s.Kühn .& Romagn .5 8 Trametes semisupina (B.& C. )Ryv .5 3 Tremella mesenterica Retz.G 9 Tricholoma aestuans Fr.5 3 T. columbetta (Fr.)Kumm .5 7 131 T. portentosum (Fr.)Quel .5 7 T. saponaceum (Fr.)Kumm .G l T. virçatum (Fr.e xFr. )Kumm .5 7 Tricholomopsis rut il ans (Schaeff.e xFr. )Sing .G 7 Tubaria conspersa (Pers.e xFr. )Fay .5 3 T. furfuracea (Pers.e xFr. )Gill .G 2 Tylopilus felleus (Bull,e xFr. )Karst .G 8 Typhula erythropus Fr.61 ,8 7 T. phacorrhiza Fr.61 ,77 ,78 ,8 7 T. giiisquiliaris (Fr.e xFr. )Henn .5 4 Tyromyces caesius (Schrad.pe rFr. )Murr .59 ,6 0 T. chioneus (Fr.pe rFr. )Karst .58 ,77 ,8 0 T. floriformis (Quél.ap .Bres. )Bond .& S .5 3 T. lacteus (Fr.)Murr .G 9 T. subcaesius David5 4 Xerocomus badius (Fr.)Kühn ,e xGilb .62 ,8 1 X. chrysenteron (Bull,e xSt.-Am. )Quél .G 2 X. parasiticus (Bull,e xFr. )Quél .60 ,9 2 X. subtomentosus (L.e xFr. )Quél .G 2 Xylaria hypoxylon (L.e xFr. )Grev .5 8 X. polymorphs (Pers.e xFr. )Grev .G 2 CURRICULUM VITAE Anna ElisatbethJanse nwer do p1 jun i195 0t eBred ageboren ,alwaa rzi j delager eschoo le nd eHBS- Bbezocht .I n196 7vin gzi jhaa rstudi ebiologi e aan,aa nd eRijksuniversitei tt eLeiden .He tkandidaatsexame nB Iwer dafge legd op 1jun i1971 .Voo rhe t doctoraal examenwerde n devolgend eonder werpend evolgend eonderwerpe nbewerkt .O phe tgebie dva nd eplantengeogra - fie, eenonderzoe k naard egren stusse nhe tFluviatie le nhe tKempen sdis trict, o.l.v. drs.J .Mennema , aanhe tRijksherbariu m teLeiden .O p het gebied vand echemotaxonomi enaa r iridoide glucosidenbi jiamzuai-soorten , o.l.v.prof.dr .R .Hegnauer , aanhe tLaboratoriu mvoo rExperimentel ePlan - tensystematiekt eLeiden .O phe tgebie dva nd etaxonomisch emycologie ,same n metM.E .Noordeloos ,ee nonderzoe knaa rd esystematie ke nfloristie kva nd e genera Marasmius, Marasmiellus,Micromphal e en Crinipellis inNederland , o.l.v.dr .C .Bas ,aa nhe tRijksherbariu m teLeiden .Va n1 septembe r197 0 tot1 septembe r197 5wa szi jal sstudentassisten tbi jhe tLaboratoriu mvoo r Experimentele Plantensystematiek in dienst bij de Rijksuniversiteit te Leiden. Na het doctoraal examen,da to p 25novembe r197 5wer d afgelegd, deed zij een onderzoek naar de systematiek en floristiekva nhe tgenu s Collybia inNederland .Va naugustu s197 6to tapri l198 0wa szi jal spromo tie-assistent indiens tbi j de Landbouwhogeschool teWageninge nvoo r het bewerkenva nee nproefschrif tove rd epaddestoele ni nenkel etype neikenbo s inDrenthe .Sind smaar t198 1i szi jopnieu wi ndiens tbi jd eLandbouwhoge school (part-time)o mee nverder ebijdrag et egeve naa nhe tmycosociologisc h onderzoekva nhe tBiologisc hStation . OH CL, MMH IH > > nMl,SHr-i3 • , Q,i-H U'MHO 0,'~l