INFORMATION FRÅN
LÄNSSTYRELSEN HALLAND
Diversity of saproxylic fungi ondecaying beech wood in protectedforests in the county of Halland
[Vedlevande svampar på bok i skyddade skogar i Hallands län]
Jacob Heilmann-Clausen
Frontpage: Hericium coralloides (koralltaggsvamp) inhabiting a large beech snag in Mårås. Theblack conks show that Fomes fomentarius (fnösketicka) have been present in an earlier stage ofdecomposition. In Halland this is consistently the case for trees inhabited by Hericium coralloides.Photo: Jacob Heilmann-Clausen.
Backcover: Jacob Heilmann-Clausen in action, Rågetaåsen nature reserve. Unknown polyporeTyromyces sp. growing on beech log at Skrockeberg nature reserve. Photos: Örjan Fritz.
Örjan Fritz edited this report, March 2005.
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Halland 2004
Contents
Contents .............................................................................................................................1Svensk sammanfattning .....................................................................................................2English summary ...............................................................................................................3Introduction .......................................................................................................................4Materials and methods.......................................................................................................6
Study area......................................................................................................................6Fungal inventory ...........................................................................................................6Environmental variables ...............................................................................................7Data analysis .................................................................................................................8Nomenclature ................................................................................................................9
Results and discussion .......................................................................................................9Overall diversity............................................................................................................9Species richness, red-listed species and indicators of habitat quality.........................10Species richness patterns at tree scale.........................................................................13Ordination of the Halland dataset ...............................................................................14Joint ordination of dataset from Halland and Denmark..............................................17Saproxylic fungi on beech wood in Halland - what is special? ..................................19Conservation value of beech forests in Halland - the European perspective..............21
Tree size .................................................................................................................22Soil conditions .......................................................................................................22Climate ...................................................................................................................23Dead wood continuity ............................................................................................23Summary ................................................................................................................26
Acknowledgements .........................................................................................................26Litterature ........................................................................................................................27
Appendix 1. Description and notes on localities studied in the project ..........................29Intensive study sites .........................................................................................................29Extensive study sites........................................................................................................33
The Oskarström area ...................................................................................................33Inland area (municipality of Hylte) .............................................................................38Southern Halland.........................................................................................................39
Appendix 2. Notes on red-listed and other interesting species recorded in the project...41Appendix 3. Ordination diagrammes...............................................................................53Appendix 4. List of species recorded in Halland.............................................................55
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Saproxylic fungi
Totalt 200 döda bokar fördelade över sju gamlabokskogsbestånd undersöktes på förekomsten avfruktkroppar av svampar. Ett standardprotokoll följ-des för ifyllande av uppgifter utvecklat inom det EU-finansierade projektet Nat-Man (www.flec.kvl.dk/natman). För varje bok antecknades alla arter av stor-svampar, utom barkväxande basidiesvampar ochnågra få ytterligare grupper, under tre olika tillfällenunder svampsäsongen 2004, baserat på förekomstenav fruktkroppar. Protokollet har tidigare tillämpats iDanmark, Nederländerna, Belgien, Slovenien ochUngern och möjliggjort en jämförelse av storsvamp-samhällen på död bokved inom Europa. Som tilläggtill nämnda intensivstudie inven-terades ytterligare 13 gamla bok-skogar mera extensivt på innehållav vedlevande svampar.
Totalt 284 arter av svampar på-träffades i undersökningen, varav20 var rödlistade. Fem arter rap-porteras här för första gångenfrån Sverige; Galerina pallida, G.pseudomniopila, G. vexans,Hohenbuehelium auriscalpiumoch Psatyrella larga. På boklågori Halland hittades i medeltal 13,1arter av storsvampar, beroende påträdstorlek, nedbrytningsstadiumoch lokal. Artdiversiteten varhögst på träd i mellanliggandenedbrytningsstadier och ökadegenerellt med trädstorleken. Denmest betydelsefulla samhällsgra-dienten, dvs det största skiftet i artuppsättning, be-fanns vara kopplad till nedbrytningsstadium av destuderade träden.
Olikheter i artuppsättningar mellan de sju intensiv-studerade områdena var begränsade. Detta kontras-terar till situationen i de flesta andra Nat-Man län-derna, där en distinkt omsättning i artuppsättningmellan de studerade områdena rapporterats i de flestafallen. Den tydligaste orsaken för denna brist på art-skillnader i Halland är den begränsade variationen iklimat och jordmån mellan de studerade lokalerna.På liknande sätt är olikheter orsakade av skillnader imarkanvändningshistoria mellan de studerade loka-lerna i Halland inte tydligt uttryckta i den totala art-stocken.
Den funna artstocken av svamp på bokved i Hallandvar mycket lik den som rapporterats från centrala
Jylland i Danmark. Båda områdena karaktäriseras avnäringsfattiga jordar och ett tämligen fuktigt ochkyligt klimat, vilka förefaller påverka svampfloransartsammansättning. Även från ett europeiskt perspek-tiv tycks klimatiska skillnader ha en mycket tydligpåverkan på svampflorans samhällsstruktur. Sålundaär svampfloran på bokved i Slovenien och Hallandöverraskande lika med tanke på det geografiska av-ståndet. Båda områdena karaktäriseras av en hög års-nederbörd, som tycks ha en tydlig inverkan på kon-kurrensförhållanden och sålunda artsammansätt-ningen på död ved.
Sett utifrån ett svenskt natur-skyddsperspektiv är bokskog-arna i Halland mycket viktigaför skyddet av vedlevandesvampar, men endast måttligtviktiga sett från en europeiskhorisont. Fyra arter medskyddsintressen rapporteradesi en högre frekvens i Hallandjämfört med andra bokskogs-regioner studerade i Nat-Man-projektet; skinntagging Denti-pellis fragilis, koralltaggsvampHericium coralloides, vedlav-klubba Multiclavula mucidaoch borstskölding Pluteusumbrosus. Dessutom är tvåtickor, Skeletocutis vulgarisoch Tyromyces sp., unika förHalland bland de bokskogs-regioner som studerats hittills.
De största skyddsvärdena noterades för lokalernaBiskopstorp, Valaklitt och Mårås samt för ett antallokaler samlade strax norr och öster om Oskarströmi den norra delen av Halmstads kommun. Substrat-förekomst, särskilt förekomsten av gamla stora trädsamt dödvedkontinuitet från en lokal till landskapsskala, identifierades som viktiga faktorer som påverk-ar det nuvarande skyddsvärdet av enskilda studer-ade lokaler. I ett europeiskt perspektiv tycks det mått-liga skyddsvärdet av vedlevande svampsamhällen ihalländska bokskogar spegla den mellanliggande niv-ån av dödvedkontinuitet på en landskapsskala, lika-väl som de halländska bokskogarnas position i ut-kanten av bokens utbredningsområde, vilket begrän-sar förekomsten av flera sällsynta vedlevande svamp-arter som lever på bokved i andra delar av Europa.
Svensk sammanfattning
Pholiota aurivellius. Photo: ÖrjanFritz.
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Halland 2004
In total 200 dead beech trees distributed across sevenold-grown beech forests stand in Halland wereinvestigated for fungal sporocarps. A standardEuropean protocol developed in the EU fundedproject Nat-Man (http://www.flec.kvl.dk/natman)was followed. On each tree all species of macrofungi,except corticoid basidiomycetes and a few othergroups were recorded on three occasions during thefungal season, based on the occurrence of sporocarps.The protocol has earlier been applied in Denmark,The Netherlands, Belgium, Slovenia and Hungary,making a comparison of macrofungal communitieson dead beech wood possible at the European scale.In addition to theintensive study sites,13 old-grown beechforests were investi-gated for saproxylic(wood-inhabiting)fungi using a moreextensive approach.
A total of 284 fungalspecies were record-ed in the project,including 20 red-listed species. Fivespecies, viz. Galeri-na pallida, G.pseudomniophila, G. vexans, Hohenbueheliumauriscalpium and Psathyrella larga are reported forthe first time in Sweden. Fallen beech trees in Hallandwere found to host an average of 13.1 macrofungalspecies, depending on tree size, decay stage andlocality. Species diversity was highest on trees inintermediate decay stages and increased generallywith trees size. The most important communitygradient (i.e. the biggest shift in species composition)was found to relate to the decay stage of the studytrees.
The differentiation in species composition among theseven intensive study sites was limited. This contraststhe situation in most of the Nat-Man countries, wherea distinct turnover in species composition amongstudy sites has been reported in most cases. The mostobvious reason for this lack of differentiation inHalland is the limited variation in climatic conditionsand soil types among the studied localities. Likewise,differences caused by variations in managementhistory among study sites in Halland are apparentlynot clearly expressed in the overall speciescomposition.
The fungal species composition on beech wood inHalland was very similar to that reported fromCentral Jutland, Denmark. Both areas arecharacterized by nutrient poor soils and a rather wetand cool climate, which both seems to affect thefungal species composition. Also at the Europeanscale, climatic differences appear to have a verypronounced impact on the fungal communitystructure. Thus, the funga on beech wood in Sloveniaand Halland appear to be surprisingly similarconsidering the geographical distance. Both areas arecharacterized by a high annual precipitation whichseems to have a marked influence on the competitive
environment and hence thespecies composition in deadwood.
In a conservation perspectivethe beech forests of Halland areconcluded to be highlyimportant for conservation ofsaproxylic fungi in a Swedishcontext and of moderateimportance in a Europeanperspective. Four species ofconservation interest, viz.Dentipellis fragilis, Hericiumcoralloides, Multiclavulamucida and Pluteus umbrosus
was recorded with higher frequency in Halland thanin other beech forest regions surveyed in the Nat-man project. In addition, two polypores, viz.Skeletocutis vulgaris and Tyromyces sp. are uniqueto Halland, among the beech forest regions studiedintensively so far. The biggest conservation interestswere recorded in Biskorpstorp, Valaklitt, Mårås andfor a number of sites aggregated north and east ofOskarström. Substrate abundance, especially thepresence of old, large diameter trees, and dead woodcontinuity at the local to landscape scale wereidentified as important factors influencing the presentconservation value of individual study sites. At theEuropean scale the moderate conservation value ofsaproxylic communities in Halland beech forestsappear to reflect the intermediate level of dead woodcontinuity at landscape scale, as well as the marginalposition of the Halland beech forests, which restrictsthe occurrence of several rare fungi inhabiting beechwood in other parts of Europe.
English summary
Trametes hirsuta. Photo: Örjan Fritz.
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IntroductionThe county of Halland hosts one of the northernmostcoherent beech (Fagus sylvatica) populations inEurope (Bengtsson 1999b). Beech established in thearea about 2000 years ago, and became an importanttree species in the mixed mainly deciduous forestsof Halland north to the Åkulla-area close to Varbergduring the subsequent 1000 years. The spread ofbeech was, like in other parts of south Scandinavia,closely linked to anthropogenic disturbances(Lindbladh et al. 2000). Beech was favoured bytemporal clearings of forests to farmland and becamean important tree species especially in the“mellanbygden” zone i.e. the transition zone betweenthe coastal lowlands dominated by open farmlandand the inland forest tracts (Bengtsson 1999b).
The present forest cover of the mellanbygden zoneexceeds 70 %, but beech is not dominant. Norwayspruce has expanded much in Halland over the lastcenturies, mostly due to plantations, and remainingbeech forests occur mostly as isolated standssurrounded by extensive coniferous forestlandscapes. Concentrations of remaining beechforests are found on the northern slopes ofHallandsåsen, at Biskopstorp NE of Halmstad andat Åkulla SE of Varberg (Fritz & Larson 2000). Evenin these landscapes, spruce forests are howeverdominating (Bengtsson 1999b; Fritz & Larson 2000;Fritz & Berlin 2002), with beech forests occurringas a mosaic of more or less confluent patches in amatrix dominated by coniferous forests or in partopen land (Hallandsåsen).
Diversity of saproxylic fungi on decaying beech wood in pro-tected forests in the county of Halland
The discomycete Bulgaria inquinans (limsvamp) is common on newly fallen beech logs and larger fallenbranches in Halland. Sporocarps are typically formed in abundance already the first year after a tree hasbroken down, but fruiting ceases already one or two years later. The species is probably present in anendophytic, symptom-less stage in living trees and initiates growth and wood decay soon after a tree ortree part has died. Photo: Örjan Fritz.
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Halland 2004
Since their establishment, the beech forests ofHalland have been used intensively for livestockgrazing, firewood collection and timber production.Some areas have even been cleared for agriculturefor shorter or longer periods (Bengtsson 1999b).None of the beech forests in Halland are accordinglypristine virgin forests, but should be seen asseminatural habitats developed in concert withcultural development. Not all types of human forestuse have been detrimental to forest biodiversity, andespecially forest grazing may have benefited speciesdepending on large trees growing in warm, semi-openforest types (Bengtsson 1999b). Dead wood hasprobably been a sparse resource in Halland beech
forests at most places and times. The most remoteand inaccessible areas (e.g. stands rich in steep cliffs,ravines etc) have probably developed more naturalstands with minimum human intervention and acontinuous presence of standing and lying deadwood. Even periods of human disaster, e.g. cattleplague, war and major epidemics, have caused largefluctuations in forest use and naturality.
In a European perspective the natural conditions ofHalland beech forests can be characterized as humid,cool and nutrient poor. The nutrient poor soil limitsthe growth rate of beech in most sites. Trunks with adiameter exceeding 80 cm are exceptional, while
Figure 1. Map showing the location of study sites inventoried for wood inhabiting fungi during theproject. Of the extensive localities Djupeåsen, Ry S, Kroksjön NÖ, Almeberget and Klinta Hallar wereonly visited at the summer visit, at which agarics were very sparsely represented by sporocarps. Back-ground photo from the beech forest reserve Råmebo. Photo: Örjan Fritz.
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Saproxylic fungi
trees with a diameter well below 50 cm may be morethan 250 years old (Bengtsson 1999a). As a conse-quence of poor growth conditions and formermanagement many beech stands are rather light openbut humid, due to the high precipitation and thespatial continuity of forests. This creates a perfectenvironment for epiphytic lichens and bryophytesassociated with beech. Halland is maybe the mostvaluable region in the whole of North Europe forthis group of organisms (cf. Fritz 2001).
Investigations of saproxylic insects have shown thatsome of the beech forests in Halland host richcommunities of rare and endangered species (Jansson2004). Preliminary investigations, especially in theBiskopstorp area, have indicated that the same maybe the case for saproxylic fungi (Fritz 2004a).Detailed investigations on the funga of Halland arehowever sparse, and very few reports have beenpublished, giving any insight in the mycologicalconservation value of Halland beech forests.
The aim of this report is to provide insight in thediversity and conservation value of saproxylic fungalcommunities in Halland; both in a regional and aninternational perspective. Recently published reports(Ódor et al 2004a, b) of the EU-funded project Nat-Man (http://www.flec.kvl.dk/natman) have given aunique opportunity to put Halland on the Europeanbeech forest map. This opportunity was taken in thecurrent project in which the standard samplingprotocol of the Nat-Man project was applied in adetailed investigation of macrofungal diversity of 200decaying beech trees in Halland. The 200 trees weredistributed in seven study sites in Dömestorp,Biskopstorp and Åkulla. In addition 13 old-grownbeech stands in other parts of Halland wereinvestigated for saproxylic fungi, using a moreextensive approach.
Materials and methods
Study areaThe county of Halland is situated in southernmostSweden along the coast of Kattegat. The climate ofthe area is temperate, with an annual meantemperature ranging from 5-7° C, and an annual meanprecipitation of 1000-1500 mm (SMHI: http://www.smhi.se). Phytogeographically the area belongsto the nemoral zone, except for a few extensive inlandstudy sites which are situated in the boreo-nemoralzone (cf. Bengtsson 1999b). The bedrock is in thewhole area dominated by acidic granites. Glacial tillsof variable thickness are present in all localities, butare often shallow or lacking on higher points. Thebedrock is hence exposed to a variable degree in allsites. The soils are generally nutrient poor and acidicwith a vegetation dominated by low grasses and dwarfscrubs (e.g. Deschampsia spp, Myrtillus spp.).Dömestorp forms an exception due to the presenceof base-rich tills supporting a relatively rich flora.Even some of the extensive inland localities havericher soils due to local occurrence of base-richamfibolit.
In all 20 localities were investigated in the project(Fig. 1). Seven of these are intensive localities inwhich a variable number of dead, fallen beech treeswere studied intensively at three occasions duringthe fungal season. Another 13 sites were extensivelocalities, visited only at one occasion, at which thewhole locality were investigated more extensively.All study sites are at present unmanaged, but to avariable degree marked by former management.
Fungal inventoryFor the intensive study a total of 200 dead beech trees,distributed across seven individual localities wereselected. At each locality trees were selected withthe intention to get a comparable and balancedrepresentation of predefined tree decay and sizeclasses across all localities. In practice trees were
Table 1. Overview of intensive localities in Halland giving basal data on forest history, numbers and sizesof study trees.
*Estimates of last year since cutting respectively max tree age.
Locality Number of study
trees Min, average and max
DBH of study trees Years since last
cutting Max tree age Dömestorp 50 20 - 48.5 - 68 cm 20* 200 Holkåsen 50 33 - 47.2 - 70 cm 50* 300 N Kroksjön 25 34 - 54.2 - 92 cm 12 250 Kvinnsåsen 15 31 - 42.8 - 50 cm 30* 270 Trälhultet 10 42 - 57.1 - 78 cm 12 270 Valaklitt 25 23 - 52.5 - 68 cm 55* 250* Äskemossen 25 20 - 42.6 - 56 cm 30* 200*
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Halland 2004
selected by walking across each locality filling up apredefined matrix until all size/decay classcombinations were equally represented. Trees, witha standing snag higher than c. 8 m, were not consid-ered relevant for the project and were hence omittedfrom the selection procedure. The variation innumbers of selected trees at each site does partlyreflect the availability of fallen trees but was decidedbefore the project start in dialog with Örjan Fritz. Itshould be noted that it was not possible to fill out thesize/decay class matrix in the same way in all sitesdue to natural and management related differencesin tree sizes. Accordingly the average size of studytrees differ considerably among sites (cf. Table 1).
In the intensive study sitesfungal inventories wereconducted in late summer(12-18 Aug), mid-autumn(29 Sept – 7 Oct) and lateautumn (1-4 Nov). At eachoccasion all selected trees,including eventual snag andbranches thicker than 10 cm,were carefully inventoriedfor fungal sporocarps, whichwere either identified in thefield or collected for micro-scopical identification.Within the basiodiomycetesall morphological groups,ex-cluding fully resupinatecorticoid fungi, were in-cluded, while non-stroma-tic pyrenomycetes and in-oper-culate disco-myceteswith sporocarps regularlysmaller than 10 mm wereexcluded from the asco-mycetes.
Extensive localities werevisited only once, at midautumn (1-9 Oct), and sur-veyed by walking around in-specting all suitable piecesof dead beech wood forfungal sporocarps. One tothree hours of field workwas used on each extensivelocality, mainly depending
on size and dead wood abundance. For logisticreasons some localities were however investigatedmore briefly, see appendix1 for details.
Environmental variablesEnvironmental variables were recorded at both thetree level and the site level. Tree variables includetree type (uprooted, broken at root neck, broken withdistinct snag), dbh (diameter at breast height), decaystage, moss cover, soil humidity, wind exposure andsun exposure. In addition the relative cover/distribution of two key wood decayers, Fomesfomentarius and Kretzschmaria deusta, were noted.
Site variables include geographical location (eastcoordinate, north coordinate, northeast coordinate,northwest coordinate), climatic key characteristics,soil type, maximum tree age and time since lastcutting. For details see Table 2.
Fomes fomentarius (fnösketicka) is clearly themost important decay agent in beech forests inHalland. It causes a vigorous white rot in infectedwood. Occupied logs are often almost completelydecayed in 10 years. Photo: Örjan Fritz.
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Table 2. List of recorded tree and site variables used in the analysis of the datasets. Site variables, writtenin italics, were only included in the analysis of the joint Halland/Danish dataset.
* For further details see Heilmann-Clausen and Christensen 2003, **Frich et al., 1997, Laursen et al.,1999, http://www.smhi.se, ***Bengtsson 1999a, Fritz & Berlin 2002, Niklasson 2003, Heilmann-Clausen & Christensen 2005 and Örjan Fritz (pers. comm.).
Data analysisApart from simple numerical comparisons a numberof more advanced analytical tools were used toevaluate the properties of the fungal dataset and tocompare it with the Nat-Man datasets from otherEuropean countries.
Detrended corresponden-ce analysis (DCA) (Økland1990) was used to investigate the overall structureof the dataset(s). DCA is an indirect ordinationtechnique, which means that the dataset is analysedbased solely on the presence of species acrosssamples (in this case the study trees). DCA order
Variable name scale Range Details DBH cm 20 - 92 Measurement (1.3 m above ground)
Decay stage Class (ordinal) 1 - 6 From undecayed (stage 1) to completely decayed (stage 6)*
Bark cover % 0 - 100 Estimated to the nearest 10 % of the log surface
Soil contact % 1 - 100 Estimated to the nearest 10 % of the log surface
Soil humidity Class (ordinal) 1 - 5 From dry soils (stage 1) to partly inundated soils (stage 5)*
Wind exposure Class (ordinal) 1 - 5 From open hilltops and edges (stage 1) to sheltered hollows and ravines (stage 5)*
Sun exposure Class (ordinal) 1 - 5 From fully exposed openings (stage 1) to closed, multi-layered forest (stage 5)*
Moss cover % 0 - 100 Estimated to the nearest 10 % of the log surface
Tree Type Class (nominal) Four types distinguished (Uprooted, Broken at root neck, Broken with distinct snag, Rebroken)
Fomes cover Class (ordinal) 0 - 3 From absent (stage 0) to dominant on wood surface(stage 3)
Kretszchmaria cover Class (ordinal) 0 - 3
From absent (stage 0) to dominant on wood surface(stage 3)
Site Class (nominal) One variable stage for each site
Region Class (nominal) Seven regions distinguished (Møn, C Zealand, N Zea-land, C Jutland, S Halland, C Halland and N Halland)
East coordinate km 33 - 246 Based on standard maps
North coordinate km 11 - 247 Based on standard maps
Northeast coordinate km 121 - 333 Based on standard maps
Northwest coordinate km 0 - 225 Based on standard maps
Soil/bedrock type Class (nominal) Five types distinguished (granite, sand, sandy till, loamy till and lime stone)
Average Jan. Temperature ºC -1.5 to 0.8 Based on data from DMI and SMHI**
Average July temperature ºC 15.5 - 16.7 Based on data from DMI and SMHI**
Annual average temperature ºC 6.5 - 8.1 Based on data from DMI and SMHI**
Average annual precipitation mm 586 - 1150 Based on data from DMI and SMHI**
Years since last cutting Years 10 - 80 Based on published and unpublished data***
Max tree age Years 140 - 350 Based on published and unpublished data***
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Halland 2004
samples in the so-called ordination space based onsimilarities in species composition. The ordinationspace is defined by ordination axes which expresscommunity gradients in the dataset. The firstordination axis in DCA is always the strongestcommunity gradient i.e. the gradient which mostefficiently expresses similarities and dissimilaritiesin species composition among samples. Subordinaryaxes express remaining variation not expressed bythe preceding axes. DCA ordination axes areassociated with two numeric descriptors: The Eigen-value express how well the axis expresses variationin the dataset while the gradient length is a measureof species turnover (á-diversity) along the axis.
Kendall rank correlation (Legendre & Legendre1998) was used to evaluate relations between fungalspecies richness per tree and investigatedenvironmental variables. Rank correlation waspreferred from its parametric alternatives due to itsinsensitivity to non-normal data. Kendall rankcorrelation was also used in the interpretation ofcommunity gradient extracted by DCA ordination.
One way ANOVA (Zar 1999) was used as analternative to Kendall rank correlation for testingrelations between species richness (or ordination axisscores) and variables on nominal scale (e.g. tree andsoil type)
Multiple linear regression (Zar 1999) was used toevaluate complex relations between species richnessscores and several independent variables at one time.Selection of variables for the model was mademanually, guided by correlation analyses and one wayANOVA’s.
NomenclatureNomenclature in general follows Hansen & Knudsen(1990, 1992, 1997).
Results and discussion
Overall diversityIn total 284 fungal species were identified in theproject. Of these 244 are macrofungi according tothe Nat-Man criteria, while 40 species are corticoidbasidiomycetes or non-stromatic pyrenomycetes. Thetrue diversity within the two latter groups in the studyarea is without doubt much higher than recorded, asonly occasional collections of the most conspicuousspecies were made within these groups. Of themacrofungi, 197 species were recorded on the 200intensively studied trees, while 47 species wererecorded as supplementary on the intensive localitiesor in the surveys of extensive localities.
The most common species in the intensive datasetare shown in Fig. 2. Only Xylaria hypoxylon andFomes fomentarius were recorded on more than halfof the study trees. Six of the ten most frequent speciesare agarics. It is important to notice that the fact thata species is common not necessarily implies that it isa dominant decay agent. Fomes fomentarius and to alesser extent Eutypa spinosa are truly dominantspecies occupying large territories already in livingtrees. Stereum hirsutum and Panellus serotinus arerestricted to wood in early decay stages, but do oftenoccupy rather large wood volumes, judged bywidespread appearance of sporocarps. Galerinamarginata, Mycena haematopus and Psathyrellapiluliformis are mostly seen on decayed wood and
0%10%20%30%40%50%60%70%80%
Xylaria
hypoxy
lon
Fomes
fomen
tarius
Marasm
ius al
liace
us
Galerin
a marg
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Lacc
aria a
methys
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sa
Mycen
a haem
atopu
s
Psathy
rella
piluli
formis
Panell
us se
rotinus
Perc
enta
ge o
f tre
es
Figure 2. The ten most commonly recorded wood inhabiting macrofungi in the project. Blue columns in-dicate pyrenomycetes, green polypores, orange agarics and pink crust-forming corticoid basidiomycetes.
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Saproxylic fungi
mostly in small localized groups. It is uncertainwhether this correlates to small occupied territoriesin the decaying wood or reflect a localizedsporulation strategy. Marasmius alliaceus andXylaria hypoxylon both occur throughout most of thedecay process. Sporocarps typically occur scatteredon several parts of decaying trees, which probablyreflect the presence of several rather small, individualmycelia. Finally Laccaria amethystina is anectomycorrhizal fungus occurring mostly on stronglydecayed wood. Its role in wood decay is unknown,but it might be actively involved in the breakdownof residual wood components or withdraw nutrientsfrom mycelia of wood decay fungi (Heilmann-Clausen 2003).
At least five of the recorded species, viz. Galerinapallida, G. pseudomniophila, G. vexans, Hohen-buehelium auriscalpium and Psathyrella larga, arereported as new to Sweden, while Cereceomycessulphurinus is reported for the second time inSweden, with the previous record, from Femsjö,dating back to 1940. In total 20 of the recordedspecies are listed in the Swedish redlist (Gärdenfors2000). Two of the recorded species appear to beundescribed and may thus be new to science.
Species richness, red-listed species and indicatorsof habitat qualityThe highest species number from a single localitywas recorded in Holkåsen, where a total of 145
Figure 3. Total (columns) and red-listed (dots) species numbers recorded on each study site. Darkcolumns indicate intensive study sites. Red-listed species corresponds to Gärdenfors (2000).
Figure 4. Number of indicator species recorded on each study site. Columns refer to the Danish indicatorsystem (Heilmann-Clausen & Christensen 2000), dots to the European system (Christensen et al. 2005).Localities are listed in the same order as in Fig. 3.
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11
Halland 2004
Table 3. Overview of red-listed species, indicator species and species new to Sweden, recorded in theinvestigated localities. The column “Biskopstorp” present a summary of species recorded in all Biskops-torp localities. Yellow shading indicates that the species was not recorded on a given locality in thecurrent project.
1According to Gärdenfors (2000); VU: Vulnerable; NT: Near threatened; DD: Data deficient. 2Christensenet al. (2005). 3Heilmann-Clausen & Christensen (2000). 4Hallingbäck & Aronsson (1998), Nitare (2000).5Reported here as new to Sweden.
species was identified. The lowest number of species(37) was recorded from Ödegärdet (Fig. 3). Therecorded species richness was generally highest inthe intensive localities, and among these, recordedspecies richness generally increases with the numberof studied trees. It is however remarkable that therecorded species richness in Dömestorp, with 50
studied trees, is lower than in both N. Kroksjön andValaklitt, each with 25 studied trees. Among theextensive localities, Mårås stand out as exceptionallyspecies rich.
Numbers of red-listed species recorded per localitydoes largely reflect the recorded species richness per
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Inonotus cuticularis VU X X X 1 1 1 2Phleogena faginea VU X X 1 1 1 1 1 1 1 1 7Multiclavula mucida VU X 1 1 1 1 1 1 1 1 8Peziza saniosa VU X 1 1Volvariella bombycina NT X X X 1 1 1 2Camarops tubulina NT X X X 1 1 1 1 1 1 1 1 1 8Dentipellis fragilis NT X X X 1 1 1 1 1 1 1 1 8Hericium coralloides NT X X X 1 1 1 1 1 1 1 1 1 1 1 1 1 12Ossicaulis lignatilis NT X X X 1 1Pluteus umbrosus NT X X X 1 1 1 1 1 1 1 1 7Lentinellus vulpinus NT X X X 1 1 1 2Hypoxylon cohaerens NT X X 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 18Ceriporia excelsa NT X 1 1 1 1 1 1 1 1 1 8Lentaria epichnoa NT X 1 1Mycena renati NT X 1 1 1 1 1 1 1 7Bolbitius reticulatus NT X 1 1 1 1 1 1 1 6Stypella subgelatinosa NT X 1 1Gloeohypochnicium analogum DD X 1 1 2Ceraceomyces sulphurinus DD ? 1 1 ? 1Hohenbuehelia auriscalpium5 X X 1 1Lentinellus ursinus X 1 1 1 2Omphalina epichysium X X 1 1 1 2Pluteus luctuosus X X 1 1 1 1 1 1 1 1 1 8Pluteus pellitus X X 1 1Pluteus phlebophorus X 1 1 1 1 1 1 1 1 7Inonotus nodulosus X 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 15Eutypa spinosa X 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 24Henningsomyces candidus X 1 1 1Inonotus obliquus X 1 1 1 3Nemania atropurpurea X 1 1 1 1 1 1 1 1 1 1 1 1 1 1 13Nemania chestersii X 1 1 1 1 1 1 1 1 1 1 1 1 1 12Pholiota aurivellus X 1 1 1 1 1 1 1 1 1 1 1 1 11Stereum subtomentosum X 1 1 1 1 3Ascotremella faginea X 1 1 1 1 1 4Oxyporus corticola X 1 1 1Pluteus nanus X 1 1 1Pluteus plautus X 1 1 1 1 1 4Pluteus podospileus X 1 1 1 1 1 1 1 1 7Pluteus salicinus X 1 1 2Pluteus semibulbosus X 1 1 1 1 1 1 1 1 1 1 1 10Skeletocutis vulgaris X 1 1 1 1 1 1 1 1 1 1 1 10Stigmatolemma urceolata X 1 1 1Galerina pallida5 1 1Galerina pseudomniophila5 1 1 1 1 3Galerina vexans5 1 1 2Psathyrella larga5 1 1 1 1 3Ascocoryne sp. 1 1 1 1 1 1 1 6Tyromyces sp. 1 1 1 1 4Number of redlisted species 8 10 10 6 7 0 1 12 6 6 7 4 3 2 1 4 5 2 4 4 3 3 0 17Number of Indicators (EU) 3 4 5 1 4 0 0 7 4 2 4 3 0 0 0 3 3 1 4 2 1 1 0 10Number of Indicators (DK) 10 14 14 8 7 2 1 18 13 9 13 7 4 6 4 9 7 6 11 9 4 6 2 24Number of ”Signalarter” (S) 13 17 14 9 11 0 2 22 9 9 11 5 5 4 3 4 6 2 7 8 5 3 0 30
12
Saproxylic fungi
locality (Fig. 3). Among the extensive localitiesÖdegärdet, Frodeparken and Skubbhult stands outas comparably poor in red-listed species, whileRågetaåsen has a remarkable high number of red-listed species on record, considering the rather lowoverall species richness.
Two different indicator systems were used to evaluatethe conservation value of individual localities in thestudy (Fig. 4). In both case a few records made byÖrjan Fritz within the last years have been includedin the dataset. Of the 42 indicators proposed forvaluation of Danish beech forests (Heilmann-Clausen& Christensen 2000b) 24 was found in the presentstudy. N. Kroksjön was the high-scorer with 16indicators, closely followed by Holkåsen, Valaklittand Mårås. Based on experiences with use of theDanish indicator system in Denmark and Europe apreliminary pan-European indicator system wasproposed by Christensen et al. (2005) (see alsoHeilmann-Clausen & Christensen 2004a). Of the 21species included in this scheme, ten were recordedin Halland: Five were recorded in the intensive studyand another five as supplementary records inintensive localities or from the extensive study sites.The highest number of European indicators was
recorded from N. Kroksjön, but it should be remarkedthat two of these were not recorded in 2004. In generalthe two indicator systems result in a comparableranking of the localities (Fig. 4). Especially Öde-gärdet, Frodeparken, Blåalt and to some extentSkubbhult stands out as localities of limited valuefor fungi inhabiting beech wood according to bothindicator schemes, while Råmebo, Skrockeberg,Mårås, Valaklitt, Dömestorp and several of theBiskopstorp localities are consistent high scorers. Amore general assessment of the conservation valuefor saproxylic fungi for all study sites are given inTable 4. This assessment accounts for records of otherinteresting wood decay fungi, and even take intoaccount more general aspects, like the size of eachlocality and its future potential as judged by theauthor.
An overview of recorded red-listed species, indicatorsand other species of special interest are given in Table3. More details on individual localities are given inAppendix 1, while the conservation value of theHalland beech forest in a European perspective isdiscussed in the last paragraph of the Results anddiscussion section.
Ectomycorrhizal fungi are very common on strongly decayed wood in Halland beech forests, probablyreflecting that dead wood offers an important nutrient resource in the generally nutrient poor forest soils.Here sporocarps of the ectomycorrhizal agaric Laccaria amethystina (ametistskivling) occur closelyassociated with Mycena haematopus (blodhätta), which is a more typical wood decay fungus. Photo:Jacob Heilmann-Clausen.
13
Halland 2004
Species richness patterns at tree scaleOn the 200 intensively studied trees a total of 2619unique records were made representing a total of 197species. Species richness per tree varied from 1 to27, with an average of 13.1. All figures are close tothe averages from the Nat-Man project (Table 5).
Average species richness per tree varied from 10.7in Dömestorp to 16.0 in Valaklitt (Fig. 5) and wasstrongly correlated also with dbh and decay stage ofstudy trees as well as with their degree of Fomescover (Fig. 6, Table 6). The multiple regression modelshow that site, decay stage and dbh all contributeindividually to explain recorded differences inspecies richness between trees. Finally there is a smallbut significant effect of Kretzschmaria cover, whilean effect of Fomes cover is delimited (Table 7). Leastsquare means of the complete model show thatspecies richness per tree is significantly lower inDömestorp than in Holkåsen, Valaklitt and Kvinns-
åsen (Fig. 7), i.e. after accounting for the effects ofdbh, decay stage and Kretzschmaria cover. It isbeyond the scope of this report to try to interpret thisdifference, but it should be emphasized thataccidental differences in rainfall during the fungalseason in 2004 may be responsible for part of thedifference. The relations between species richnessper tree and the variables decay stage and dbh seemto be fairly universal for beech forests in Europe(Heilmann-Clausen & Christensen 2003, 2004b;Ódor et al. 2004a). Thus species richness increaseswith increasing tree size, which is not surprising, andpeaks on trees in intermediate stages of decay, whichmost likely reflect a peak in the number of availableniches for saproxylic fungi. See Heilmann-Clausen& Christensen (2003) for further details. The negativerelation between Kretzschmaria cover and speciesrichness, probably reflect a defensive decay strategiin Kretzschmaria deusta. The species keep occupiedwood dry in order to minimize competition from
Table 4. A general assessment of the conservation value in a Swedish context of the study sites based onthe occurrence of saproxylic fungi on beech wood, and the future development potential.
Halland Denmark Belgium The Netherlands
Hungary Slovenia Totals/ Averages
Number of study trees 200 200 195 198 207 214 1214 Number of study sites 7 6 1 8 2 2 26 Number of species 197 257 190 155 227 207 485 Number of species occurrences
2619 2938 3264 2300 2635 1819 15.575
Average number of species per tree
13.1 14.7 16.7 11.6 13.1 9.1 12.8
Table 5. Summary of species diversity data from the Nat-Man project and the intensive investigation of200 beech logs in Halland.
* According to Heilmann-Clausen & Christensen 2000; **According to Christensen et al. 2005
Conservation value Localities Criteria Very high Holkåsen
N Kroksjön Valaklitt Getabäcken Mårås Råmebo Skrockeberg
• 4-10 Red-listed species • 7-14 Danish indicator species* • 3-5 European indicator species** • Several species of conservation concern recorded in larger populations • Rather large localities with good prospects for future development
High Dömestorp Trälhultet Kvinnsåsen Äskemossen Nissaström Rågetaåsen Myskebackarna Svarta Klippan Vallåsen
• 2-8 Red-listed species • 4-10 Danish indicator species* • 1-3 European indicator species** • Species of conservation concern recorded as single individuals or in small populations • Small to large localities with moderate to good prospects for future development
Low to Moderate Blåalt Frodeparken Skubbhult Ödegärdet
• 0-3 Red-listed species • 2-6 Danish indicator species* • 0-1 European indicator species** • Species of conservation concern recorded as single individuals • Small to large localities with moderate to good prospects for future development
14
Saproxylic fungi
basidiomycetes. As a result the decay process isslowed down, and the realised actual species richnessis decreased (Heilmann-Clausen 2001).
Ordination of the Halland datasetThe dataset used in the ordination analysis contained,after exclusion of species poor trees (< 5 species),rare species (< 3 occurences) and outliers, 2459species records, representing 129 species on 179trees. The DCA ordination of this dataset revealedone very clear community gradient expressed by axis1 with an Eigen-value of 0.56 and a gradient lengthof 5.13 SD units. Eigen-values, as well as gradient-lengths, of subsequent axes are much smaller (Axis2: 0.19 and 2.74 respectively; Axis 3: 0.17 and 2.40respectively) pointing to a much lower importancefor resolving variation in the dataset. Subsequentordination axes above axis 3 were not extracted. Thedominant community gradient expressed by axis 1is clearly successional, dividing trees according totheir decay stage. The second and much weakergradient to some degree divide trees according tostudy sites, while the third gradient relates to dbh aswell as site (Table 6, Fig. 8). The optima of study
sites along axis 2 give few hints to understandingthe underlying community gradient. The lowest axisscores are generally obtained by trees in Dömestorpand Valaklitt, which are the southern-most andnorthern-most localities, respectively. Dömestorp ischaracterized by rather base-rich glacial tills, whileValaklitt are on a south facing slope with rather deepdeveloped soils. The highest axis scores are obtainedby trees in N. Kroksjön, Kvinnsåsen and Äske-mossen. The two first localities are situated in theBiskorpstorp area, while the third is in the Åkullaarea. The three sites seems not to have much incommon in general aspects and management historybut are at least partly characterized by very thin soilsand marginal growth conditions for beech. It is thuspossible that the axis express a weak soil richnessgradient.
The distribution of fungal species in the ordinationspace show that Ascocoryne sarcoides, Hypoxylonfragiforme, Inonotus nodulosus, Nectria coccinea,Schizophyllum commune and Trametes hirsuta arespecies with optima on fresh, only slightly decayedwood (Appendix 3). Strongly decayed wood, on the
Peziza micropus (kortfotad storskål) was recorded quite abundantly in several study sites, mostly on di-stinctly to strongly decayed logs and snags. For unknown reasons it was not recorded at all in Dömes-torp, while it occupied 40 % of the trees in Holkåsen, and at least 16 % of the trees in all other intensivestudy sites. Photo: Örjan Fritz.
15
Halland 2004
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Figure 7. Predicted mean species richness per tree per site based on least square means of the multipleregression model. Whiskers indicate 95% confidence intervals. Similar letters indicate localities withmeans not significantly different from each other. For details of the model see Table 7.
Figure 5. Average species richness per tree (columns) and average DBH of study trees (dots) in each ofthe intensive study sites.
Figure 6. Relations between species richness per tree and the key variables decay stage and DBH of thestudy trees. The whiskers on the left hand figures indicate standard deviations of the average speciesrichness for each decay stage.
16
Saproxylic fungi
other hand, is characterized by the occurrence ofseveral ectomycorrhizal fungi, e.g. Cantharellustubaeformis, Cortinarius anomalus, Laccaria laccataand Russula ochroleuca, but also by strictlysaproxylic late stage agarics e.g. Bolbitius reticulatusand Pluteus spp. Species with a more central positionin the ordination space either have their optima onintermediately decayed trees, or have no distinctdecay stage preference. The latter is the case for someof the most common pyrenomycetes, Eutypa spinosa,Kretzschmaria deusta and Xylaria hypoxylon whichhave a defensive decay strategy and persist fruitingthrough most of the decay process (Heilmann-Clausen 2001). Species with a preference for treesin intermediate decay stages include some corticoidand polyporoid species, e.g. Antrodiella semisupina,Polyporus varius, Steccherinum ochraceum as wellas cord forming basidiomycetes, e.g. Armillaria lutea,Hypholoma fascicularis and Lycoperdon spp. Thedistribution of species along axes 2 and 3 ads little
to the interpretation of the expressed communitygradients and is not discussed here. Considering thelow Eigen-values of both axes it is most reasonableto treat them as expressions of weak tendencies inoverall species composition relating to weak andpartly unknown habitat factors.
In summary the ordination analysis shows that themost important community gradient in the Hallanddataset relates to the decay stage of the study trees.This result is in agreement with the situation in otherEuropean countries (Heilmann-Clausen 2001; Ódoret al. 2004a). The rather limited geographicaldifferentiation between individual study-sites inHalland along subsequent axes in the ordination, onthe other hand, is in contrast to the situation in allNat-Man countries involving more than one studysite (Ódor et al. 2004b). The obvious reason for thislow degree of differentiation in Halland is that thevariation in climatic conditions and soil types among
Table 6. Overview of univariate relations between environmental variables and species richness and ordi-nation scores per tree. Site variables given in italics were only included in the analysis of the ordinationaxes resulting from the joint ordination of the datasets from Halland and Denmark. For relations betweenenvironmental variables and species richness Kendalls tô are given, except for relations involving theordinal variables decay stage, soil moisture, wind exposure, tree type, soil/bedrock type, site and regionwhere F-statistics from one-way ANOVA’s are given.
Significance levels: * P < 0.05; ** P < 0.01; *** P < 0.001; **** P < 0.0001.
Halland Halland + Denmark
Variable Species richness
DCA axis 1
DCAaxis 2
DCAaxis 3
DCA axis 1
DCAaxis 2
DCAaxis 3
DBH 0.40**** -0.21**** 0.17**** Decay Stage 4.58** 146.7**** 2.32* 2.30* 149.2**** 2.44* Bark cover 0.68**** 0.09* Moss cover -0.27**** -0.10* Soil Contact -0.55**** -0.44**** -0.20**** Sun exposure 2.44* 7.28**** Wind exposure 2.68* 31.94**** Soil humidity 4.16* 4.03* 3.72* Tree type 28.67**** 3.37* 12.39**** 34.67**** Fomes cover 0.20*** 0.15* 0.18**** -0.30* Kretzschmaria cover 0.15* -0.11* 0.25**** Site 4.26** 6.03**** 5.14**** 1.79* 52.52**** 4.65**** Region 3.41* 106.1**** 7.97**** East coordinate -0.16**** North coordinate -0.55**** Northeast coordinate -0.33**** Northwest coordinate -0.48**** Soil/bedrock type 3.89** 127.3**** 7.63**** Average Jan. Temperature 0.50**** Average July temperature -0.12* 0.55**** Annual average temperature 0.56**** Average annual precipitation -0.59**** -0.10* Years since last cutting -0.09* 0.33**** Max tree age 0.24**** -0.09*
17
Halland 2004
localities is more limited than in other Nat-Mancountries.
Joint ordination of dataset from Halland andDenmarkA further DCA ordination was performed using theHalland dataset combined with the correspondingDanish Nat-Man dataset. Again species poor samples,rare species and outlying samples were omitted. Theresulting dataset contained 5283 species records,representing 185 species on 366 trees. The DCAbased on this dataset resulted in the expression oftwo distinct community gradients, with Eigen-valuesof 0.47 and 0.30 respectively and gradient lengths of4.51 and 3.91 SD units. The third ordination axiswas considerably weaker (Eigen-value 0.17; gradientlength 3.27 SD units) and is not evaluated any furtherhere, since it was not found to relate distinctly torecorded tree or site variables.
The first DCA axis does, like in the pure Hallanddataset, represent a clear decay gradient, while thesecond axis represents a distinct geographicalgradient (Table 6). Trees from sites in Halland forma distinct cluster along axis 2, together with trees in
Silkeborg Vesterskov and Velling in Central Jutland.Another cluster is formed by trees in the two sites inZealand. Finally trees from Store Klinteskov, Mønforms a third cluster opposite to the Halland/CentralJutland cluster (Fig. 9) The distribution of sites alongAxis 2 relates to a concurrent, distinct turnover insoil types and climatic conditions (Table 6), and amultiple regression analysis revealed that both factorscontribute individually to explaining the distributionof sample trees along the axis (result not shown).From an overall perspective, the axis accordinglyexpress the shift from dry and warm, calcareous sitesin Møn, over typical mesic beech forest on richersoils in Zealand to wet and cool beech forest on poorsoils in central Jutland and Halland.
With respect to species composition it is striking thata high number of ectomycorrhizal agarics, e.g.Cantharellus tubaeformis, Cortinarius spp., Inocybepetiginosa, Laccaria spp., Lactarius spp. and Rusullaspp. have optima in Halland and Central Jutland(Appendix 3). All are common species in Denmarkand south Sweden, but were only rarely recorded ondead wood in Danish study sites on richer soils. Thisprobably reflects that dead wood is an important
Phlebia radiata (ribbgrynna) is a common, but beautiful corticoid fungus on dead beech wood in Halland.It is most common on rather hard wood (decay stage 2), and disappear gradually as decay proceeds.Photo: Örjan Fritz.
18
Saproxylic fungi
Axis 1
Axi
s 2
regionDK, MønDK, C ZealandDK, N ZealandDK, C JutlandS, S HallandS, C HallandS, N Halland
Axis 1
Axi
s 2
SiteDömestorpHolkåsenN KroksjönKvinnsåsenTrälhultetValaklittÄskemossen
Figure 9. Diagram showing the location of individual study trees in the ordination space defined by axis 1and 2 in the DCA ordination of the joint datasets from Halland and Denmark. Trees located in differentregions are indicated by different symbols.
Figure 8. Diagram showing the location of individual study trees in the ordination space defined by axis 1and 2 in the DCA ordination of the Halland dataset. Bubble sizes illustrate the decay stages of individualtrees. Large bubbles are most decayed.
19
Halland 2004
nutrient resource for ectomycorrhizal fungi mainlyon naturally nutrient poor soils. True wood decayfungi with optima in Halland e.g. Hericium corall-oides, Nemania atropurpurea, Pluteus podospileusand Skeletocutus vulgaris may, on the other hand, bebenefiting from a more cool and moist climate. Thesame is the case for the lichenized Multiclavulamucida.
A mixing of climatical and soil type aspects is evidentalso in the other end of the gradient. Thus, severalspecies with optima in the calcareous beech forestson Møn and C Zealand are known or suspected tohave a preference for base-rich soils. This is e.g. thecase for Ceriporia reticulata, Micromphale brassi-colens, Pluteus romellii and Steccherinumfimbriatum. Other species like Camarops polysper-
ma, Ischnoderma resinosum, Phellinus ferruginosusand Xylaria polymorpha which also have optima inthe Danish sites outside Jutland, probably reflect theclimatical aspect of the gradient, i.e. a preference fora drier and/or drier climate than occurring in Halland.
Saproxylic fungi on beech wood in Halland – whatis special?Of the 197 macrofungi (ss. Nat-Man) identified onthe 200 intensively studied trees in Halland, 19 werenot recorded in any of the original Nat-Man countries.Most of these were recorded on one or two trees only,while five species were recorded on eight trees ormore (Table 8). Most species overrepresented inHalland are ectomycorrhizal agarics or leaf-littersaprotrophs. This probably reflects the generalnutrient poor soils in Halland. Several, e.g. Cysto-
Eutypa spinosa
0%
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Figure 10. Frequencies (% of study trees) of four key primary decay agents in beech wood in Halland andeach of the Nat-Man countries.
Variable Degrees of freedom
Prefix of parameter estimate
Type III Sum of Squares
Mean Square F Value Pr > F
DBH 1 + 1678.6 1678.6 105.3 <0.0001 Decay stage 5 679.9 136.0 8.53 <0.0001 Site 6 452.0 75.33 4.73 0.0002 Kretzschmaria cover 1 - 75.95 75.95 4.76 0.03
Table 7. Summary of the multiple regression model (p < 0.0001; R2 = 0.50) for fungal species richnessper tree in the Halland dataset.
20
Saproxylic fungi
Species Halland Denmark Belgium The Netherlands Hungary Slovenia
Mycena cinerella L 20% 2% 0% 2% 0% 0% Antrodiella semisupina W 16% 6% 1% 7% 2% 0% Inocybe petiginosa M 15% 2% 1% 0% 0% 0% Nemania atropurpurea W 11% 3% 0% 0% 0% 2% Lactarius blennius M 9% 3% 0% 0% 0% 1% Russula mairei M 9% 4% 1% 0% 0% 0% Cantharellus tubaeformis M 7% 1% 0% 0% 0% 1% Helvella lacunosa M? 7% 1% 0% 0% 0% 0% Hydnum rufescens M 6% 0% 0% 0% 0% 0% Mycena hiemalis W 6% 2% 1% 1% 0% 2% Ascocoryne sp W 5% 0% 0% 0% 0% 0% Skeletocutis vulgaris W 5% 0% 0% 0% 0% 0% Cystoderma amianthinum L 4% 0% 0% 0% 0% 0% Mycena metata L 4% 0% 0% 0% 0% 0% Stropharia hornemannii L 4% 0% 0% 0% 0% 0% Lepiota felina L 3% 0% 1% 0% 0% 0% Cortinarius anomalus M 3% 0% 1% 0% 0% 0% Galerina mniophila s.l. B 3% 0% 0% 0% 0% 1%
Table 8. Species overrepresented in Halland, i.e. species recorded on at least 5 trees in Halland, and atleast twice as frequent here than in any of the Nat-Man countries. Percentages refer to the fraction ofintensively investigated trees presenting the actual species in each of the countries.
Ecological codes: B: Bryophyte-associated; L: Litter saprotroph; M: Ectomycorrhizal; W: Wood sapro-troph
Species Halland Denmark Belgium The Netherlands Hungary Slovenia
Mycena arcangeliana W 0% 0% 2% 13% 51% 14% Meripilus giganteus W 0% 6% 9% 16% 3% 0% Pluteus nanus L 0% 1% 14% 0% 14% <1% Postia tephroleuca W 0% 0% 8% 16% 3% <1% Ceriporiopsis gilvescens W 0% 0% 11% 2% 6% 6%Pluteus hispidulus W 0% 1% 17% 2% 0% 2%Skeletocutis nivea W 0% 2% 3% 11% <1% <1% Xylaria polymorpha W 1% 8% 18% 4% 9% 9%Hyphodontia paradoxa W 1% 7% 0% 17% 9% <1% Pluteus salicinus W 1% 6% 5% 2% 3% 0% Stereum subtomentosum W 1% 5% 19% 27% 2% <1% Simocybe centunculus W 1% 4% 7% 0% 14% 9%Ramaria stricta W 1% 14% 1% 1% <1% 0% Mycena crocata W 2% 28% 6% 0% 32% 10% Mycena pura L 2% 1% 0% 1% 11% 2%
Table 9: List of non-Halland species, i.e. species absent or occurring with very low frequency (<5 trees)in Halland, but recorded as common (at least 10 % of trees) in one or more of the Nat-Man countries.Percentages refer to the fraction of intensively investigated trees presenting the actual species in each ofthe countries.
Ecological codes: L: Litter saprotroph; W: Wood saprotroph.
derma amianthinum, Lepiota felina, Mycenacinerella and Stropharia hornemannii are moretypical for other habitat types than beech forest,including coniferous forests, birch forests and heaths(cf. Hansen & Knudsen 1992), and are common inthese habitat types in at least some of the Nat-mancountries. Three species, viz. Antrodiella semisupina,Nemania atropurpurea and Skeletocutis vulgaris are,however, distinctly wood associated, and could be
regarded as Hallandic specialities in a Europeanbeech forest perspective. The same may be the casefor Ascocoryne sp., but it is possible that this speciesis included in the concept of other Ascocoryne speciesin some of the Nat-Man countries.
As well as typical Halland species it is worthwhileto consider distinct non-Halland species; these arelisted in Table 9. Of the listed species Mycena
21
Halland 2004
arcangeliana has distinct optima in Hungary andSlovenia, Stereum subtomentosum seems to prefersites in Belgium and the Netherlands, whileCeriporiopsis gilvescens appear to be a distinctsouthern species, lacking in both Denmark andSweden (the species is actually recorded in bothcountries, but only from very few sites). Meripilusgiganteus and Xylaria polymorpha are quite frequentin most or all of the Nat-man countries and could beconsidered as the most distinct non-Halland species,probably for climatical reasons. Mycena crocata is(almost) lacking in both Sweden and the Netherlandsand even very infrequent in Belgium. This most likelyreflects a preference for rich soils, though climaticalfactors may contribute as well.
Two very common pyrenomycetes, Eutypa spinosaand Kretzschmaria deusta were recorded with lowerfrequency in Halland and Slovenia than in the otherNat-Man countries (Fig. 10). Both are well adaptedto low water potentials (Heilmann-Clausen 2001) andare likely to be less competitive in a humid climate.Ganoderma lipsiensis was, for unknown reasons,recorded with low frequency in Halland, Denmarkand Slovenia, while Fomes fomentarius was veryfrequent in all countries, except Belgium and to alesser extend the Netherlands. The species is clearlythe most important primary decay agent in Hallandand was generally found to occupy larger volumesof wood in individual trees, than in other Nat-Mancountries. Unlike Ganoderma spp. and Kretzsch-maria deusta it causes predominantly decay in thecentral part of the trunk of living trees, making themliable to break 5-8 meter above the ground. Theresulting snags are a very prominent feature in beechforests in Halland (Bengtsson 1999b) and seem tosupport special fungal communities. Thus, Camaropstubulina, Nemania atropurpurea, N. chestersii, Sisto-trema sernanderi and Trechispora nivea were pre-dominantly found in association with standing orfallen beech snags decayed by Fomes fomentarius.The comparison of Bray-Curtis similarity indexscores shows that the overall species composition inthe Halland-dataset has most similarities to theDanish Nat-Man dataset (Table 10). While this is notsurprising, it is remarkable that the Slovenian datasetis the second most similar, though it is geographicallythe most distant. The sites in Slovenia and Hallandhave two important characteristic in common, whichare not shared by any other Nat-Man study sites:Winters are cool (average temperature of January c.-1º C) and the average annual precipitation isexceeding 1000 mm (cf. Ódor et al. 2004a). Soilconditions are entirely different between the studyareas, as the Slovenian sites are on limestone.
Therefore the climatical similarities are the mostprobable explanation for the found similarities. Threespecies, Multiclavula mucida, Galerina mniophilas.l. and Mycena pseudocorticola, were recordedfrequently in the study sites in Halland and Slovenia,but not or very scarcely in the other Nat-Mancountries. They are not important wood decayers, butall are likely to prefer a humid forest climate. Thefirst due to its algal symbiont, the second due to itsdependence on bryophyte-covered logs, and the thirdbecause of its very small, desiccation sensitivesporocarps. The Hungarian dataset is the mostdifferent from the Halland dataset according tosimilarity index scores. Also here climatical factorsare a likely explanation. Thus the Hungarian siteshave the most continental climate in the wholedataset, with large differences between summer andwinter temperatures (difference between averageJanuary and July temperatures ˜ 20º C), and a ratherlow annual average precipitation (<900 mm) (cf.Ódor et al. 2004a).
Conservation value of beech forests in Halland -the European perspectiveIn the European perspective the number of indicatorspecies (ss. Christensen et al. 2005) recorded inHalland beech forests is not too impressing (Table11). Even when all Biskopstorp localities are mergedto one area, corresponding to the situation for manyof the European sites in the table, the number ofindicators only adds up to seven (Table 12). Thisplaces the Biskopstorp area well below the mostvaluable sites in Denmark, France, UK and EasternEurope, but on line with the most valuable sitesknown so far in Skåne, i.e. Maltesholm, Ivø andSkäralid. Other investigated sites in Halland had onlyup to four indicator species in total (see Table 3).
In Fig. 11 the presence of European beech forestindicators (Christensen et al. 2005) is comparedamong the standardised datasets from Halland andthe Nat-Man countries. It is evident also in thiscomparison that the number of indicator species inHalland is in the low range, +/- equal to the situationin Belgium and the Netherlands, but well below thescores in Denmark, Slovenia and Hungary. However,the proportion of records made up by indicators inHalland is rather high. This reflects that someindicator species were recorded rather frequently inHalland. In fact three species, viz. Camaropstubulina, Hericium coralloides and Pluteusumbrosus, appeared to be more common in Hallandthan in any of the Nat-Man datasets.
From an overall point of view the beech forests of
22
Saproxylic fungi
Halland can thus be stated to have a moderateconservation value in the European perspective, whilethey appear to be highly valuable in a Swedishcontext. However, it is highly relevant to ask if theused indicator species indicate anything interesting,apart from the sheer presence of dead wood. Thisquestion is addressed in some detail in the followingsections considering potential causal factors behindthe rather low indicator scores in Halland.
Tree sizeTree size has often been quoted to be an importantfactor for endangered saproxylic organisms (e.g.Bader et al. 1995). Beech trees in Halland aregenerally rather small, and it could be argued thatsome fungal species are limited by the lack of suitable
large diameter trees. In Fig. 12 the recorded distri-butions on diameter classes are illustrated for the fourmost common indicator species in Halland.Camarops tubulina and Hericium coralloides bothappear to have a preference for large diameter trees,while Dentipellis fragilis and Pluteus umbrosus showvery unclear trends. The figure is based on sparsedata and does of course not tell anything aboutdiameter preferences of indicator species notrecorded in the study. In an extensive discussion onthe subject Heilmann-Clausen & Christensen(2004b), concluded tree size or rather tree age to beof importance mostly for species causing heart rot.The infection strategies of Camarops tubulina andHericium coralloides are unknown, but the latterspecies is suspected to be a selective successor ofFomes fomentarius and Inonotus spp. (Niemelä etal. 1995). Several others of the indicators are wellknown to be heart rot agents and it is possible thatsome of these are in fact limited in Halland due tothe scarcity of old or big trees. The hypothesis isimpossible to test based on the current material, butit is noteworthy that trees with a dbh exceeding 60cm are very scarce in several of the study sites inHalland, e.g. in Äskemossen and Kvinnsåsen, whichhad lower indicator scores and smaller maximum treesizes than the other intensive study sites (compareTable 1 and 3).
Soil conditionsAs previously discussed the fungal speciescomposition on decaying beech wood in Hallandseems to be distinctly influence by the dominantnutrient poor soils. The high proportion ofectomycorrhizal fungi in late decay stages is moststriking, but it is relevant to ask if soil conditions areof direct importance for any of the indicator species,and hence for the rather low indicator scores inHalland. Most of the indicators are normally asso-ciated with rather hard wood and for these a causalrelation between soil nutrient status andestablishment success / competitive ability is un-likely. Theoretically, it is possible that woodchemistry, as affected by soil nutrient status, mayaffect species in an unbalanced way, but as yet
Table 10. Bray-Curtis (qualitative Sørensen) similarity matrix for comparisons of the datasets fromHalland and the Nat-Man project. Higher scores indicate higher degree of similarity.
Sweden Denmark Slovenia The Netherlands Belgium Hungary
Sweden 1 0.67 0.55 0.52 0.47 0.44 Denmark 0.67 1 0.54 0.56 0.58 0.52 Slovenia 0.55 0.54 1 0.49 0.42 0.54 The Netherlands 0.52 0.56 0.49 1 0.60 0.41 Belgium 0.47 0.58 0.42 0.60 1 0.39 Hungary 0.44 0.52 0.54 0.41 0.39 1
Species
Known from Halland
Known from Sweden
Aurantioporus alborubescens X* Camarops tubulina X X Ceriporiopsis gilvescens X Ceriporiopsis pannocincta X Climacodon septentrionalis X Dentipellis fragilis X X Flammulaster limulatus X Flammulaster muricatus X Ganoderma pfeifferi X Hericium coralloides X X Hericium erinaceum X Hohenbuehelia auriscalpium X* X* Inonotus cuticularis X X Ischnoderma resinosum X Lentinellus ursinus X X Lentinellus vulpinus X X Ossicaulis lignatilis X X Phlebia nothofagi Pholiota squarrosoides X Pluteus umbrosus X X Spongipellis delectans X
Table 11: Indicator species for assessing conser-vation value of beech forests in Europe, listed byChristensen et al. (2005), and their status inSweden and Halland.
*Not listed in Hallingbäck & Aronsson(1998).
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Halland 2004
way of arguing Flammulaster limulatus and F.muricatus are the species most likely to be limited inHalland due to the nutrient poor soils, but even forthese a direct soil effect is considered most unlikelyin the current case. Thus, Flammulaster limulatus is
recorded from at least 50 sites in Swedenaccording to Sveriges MykologiskaFörening (http://www.svampar.se/index.asp), and is not restricted to regionswith base-rich soils.
ClimateAs previously discussed climaticconditions seem to play an important rolefor the fungal species composition on deadbeech wood at the European scale. It istherefore reasonable to discuss if climaticfactors play an important role for therecorded indicator scores in Halland. Thisclearly seems to be the case. Halland is atthe extreme limit of the distribution areafor several of the European indicators.This is e.g. the case for Ceriporiopsis gil-vescens, Ganoderma pfeifferi andSpongipellis delectans which are southernspecies in Europe (Ryvarden & Gilbertson1992-3), while Climacodon septen-trionalis and Pholiota squarrosoides aresouth-eastern, continental species (ownobservations). It is likely that these specieswere never represented by stable popu-lations in Halland in the past, meaning thatthe potential maximum indicator score inHalland is somewhat lower than in centralEurope. To account for this it could berelevant to include more species in theevaluation of conservation value of beechforest sites in Halland. EspeciallyMulticlavula mucida, Skeletocutis vulga-ris and Tyromyces sp. are interesting inthis context. All are potentially ofEuropean conservation interest, and aremore or less restricted to Halland beechforests in the Nat-man context. However,more knowledge on their habitatpreferences, distribution patterns andtaxonomic status (Tyromyces sp.) is need-ed before their indicator values can beevaluated in detail.
Dead wood continuityMost of the old grown beech forest remnants inHalland are small (<10 ha), isolated, but partlyaggregated at landscape scale. This is particularlythe case in the Åkulla and Biskopstorp areas and to a
Table 12. List of beech forests of conservationvalue in Europe, based on the occurrence of 21saproxylic fungi suggested as indicators of habitatquality by Christensen et al. (2005). Swedish sitesare shaded.
relevant experiments have not been published. Forspecies living in more decayed wood, effects are morelikely, because the importance of the soil/woodinterface increases as decay proceeds, due to animalactivity, physical break down of wood and transportof nutrients through fungal mycelia. Following this
Country Site name
Score from 21 possible
Slovakia Stuzica 16 Slovakia Rozok 16 Czech Republic Zofin 15 France Fontainebleau (La Tiliaie) 15 Denmark Jægersborg Dyrehave 14 Slovakia Havesová 13 UK Wood Crates 13 Denmark Suserup Skov 12 Denmark Strødam forest reserve 12 UK Denny Wood area 12 Slovenia Rajhenav Rog 11 Hungary Öserdö 11 UK Windsor Bears Rails/Wild Boar area 11 UK Windsor Highstanding Hill 10 Denmark Store Bøgeskov 9Sweden Maltesholm, Skåne 9UK Whitley Wood 9UK Norbury Park 9Denmark Romsø 8Hungary Kekes 8Slovakia Udava 8UK Lullingstone Park 8Denmark Krenkerup Haveskov 7Denmark Åhuse 7 Denmark Stenderup 7 Sweden Ivö, Skåne 7Sweden Skäralid, Skåne 7Sweden Biskopstorp, Halland 7UK Mark Ash Inclosure 7UK Gritnam Wood 7UK Ebernoe Common 7Germany Waldhaus 7Denmark Slagslunde 6 Denmark Store Klinteskov 6 Denmark Nørreskoven 6 Slovenia Krokar 6 Poland Poland Biaskidy E 6 Belgium Kerssellaerspleyn 6 UK The Mens & Cut 6 Germany Mittelsteighütte 6
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Saproxylic fungi
lesser extent in the landscapes north and east ofOskarström. Present day dead wood amounts are con-siderable in several of the individual localities, butdead wood abundance is mostly of recent date, thoughsmall spots may represent long local dead wood con-tinuity. In the present landscape the beech forests areaggregated in matrix of extensive coniferous forests(forest cover >60 %). The forest cover has increasedduring the last centuries, but at the same time thearea with beech forests have declined considerably.Most of the remaining beech forests hence have rootsback to the more extensive deciduous forests of thepast. In a European perspective this (very grossly)points to an intermediate level of dead woodcontinuity at the landscape scale: The largecontinuous beech forests of the historical Hallandmost likely secured a continuous presence of at leastsome dead wood in the landscape, unlike the situationin most of the northwest European lowlands, whereforest was very scattered and highly overused in the
past. The other end of the gradient is represented bymountainous regions in Eastern Europe, which arestill rather rich in virgin forest remnants with verynatural forest structures and, apparently, unbrokendead wood continuity since the ice age.
Generally, saproxylic fungi have good abilities forlong range dispersal and have for this reason beenquestioned as relevant indicators of habitat continuity.Nordén & Appelqvist (2001) for instance suggestedthat “many wood-decay fungi are … probably morefavoured by a regionally rich supply of microhabitatsthan by long persistence of local forest stands”.Rolstad et al. (2002) went even further and statedthat “indicator species of forest continuity (includingsaproxylic fungi; my insertion) fail to fulfil all thecriteria of a suitable indicator”. Considering the verystrict habitat requirements in some saproxylic fungiand their general lack of precision in dispersalChristensen et al. (2005) challenged this view and
stated that “wood-inhabitingfungi can provide relevantbiodiversity information,which cannot be gathereddirectly from simple structuralindicators, e.g. volume of deadwood per ha” and stated theindicator system used here tobe “especially suitable foridentifying relatively largeareas (25-10,000 ha) possess-ing dead wood continuity”. Ascientific sound testing ofthese partly conflicting viewsare far beyond the scope of thisreport, but still a closer lookon data is worthwhile.
0
2
4
6
8
10
12
Halland
Denmark
Belgium
The N
etherlan
ds
Hungary
Slovenia
Num
ber
of E
U in
dica
tors
in d
atas
et
0%
1%1%
2%2%
3%3%
4%
Pro
port
ion
of to
tal r
ecor
ds m
ade
up b
y E
U in
dica
tors
Figure 11. Total number of indicator species (ss. Christensen et al 2005)(columns)and their proportions oftotal species records (dots) in the intensive datasets from Halland and the Nat-Man countries.
DBH
0%
2%
4%6%
8%
10%
12%14%
16%
18%
Camaropstubulina
Dentipellisfragilis
Hericium coralloides
Pluteusumbrosus
Frac
tion
of tr
ees
with
spe
cies
20-39 cm40-59 cm>60 cm
Figure 12. Frequencies of the four most common indicator species (ss.Christensen et al. 2005) on different tree size classes in the Hallanddataset.
25
Halland 2004
If rare wood-inhabiting fungi were predominantlysubstrate limited a close correlation between localdead wood quality and amounts and indicator scoreswould be expectable. As mentioned above, there infact seem to be a positive relation between the pre-sence of old, large diameter trees and high indicatorscores, pointing to a possible affect of substrate limi-tation. In other cases such a relation is not evident,while an effect of lacking dead wood continuity isprobable. Thus, no indicator species were recordedin three of the most isolated stands, viz. Blåalt,Skubbhult and Ödegärdet, though the amounts ofdead wood in all sites are considerable. This couldreflect local extinction of indicator species, due tobroken dead wood continuity locally and in thesurrounding landscapes. Sites with high indicatorsscores, on the other hand, are either large (Mårås) orsituated in landscapes rich in old-grown beech forestreserves. Here dead wood continuity, either locallyor at landscape scale, is more likely to have persistedduring the last centuries.
At the species level it is remarkable that several ofthe indicators have a limited distribution in Hallandaccording to known data. This could reflect localisedsurvival of remnant populations during a bottle-neckin dead wood abundance, like suggested for the tineid,
saproxylic moth Scardia boletella in the samelandscape (Fritz 2004b). Scardia boletella is ratherwidely distributed in Halland but the core-area isclearly in the Biskopstorp/Oskarström area. At leastfour potential fungal indicator species have a similardistribution pattern, viz. Lentinellus vulpinus, L.ursinus, Tyromyces sp. and Multiclavula mucida (forthe two latter species see Fig. 13). There is no doubtthat wood-inhabiting have better abilities for longrange dispersal than Scardia boletella, which appearto be poorly able to colonize new habitats situatedjust a few km’s away from existing ones (Fritz2004b). Scardia, however has much denserpopulations than most of the rare wood-inhabitingfungi discussed, and occur typically on dozens ofFomes-infected trees in occupied localities. Insummary, rare wood-inhabiting fungi, including theindicators, tend to have much stricter habitatrequirements but better dispersal ability than Scardiaboletella and probably many other saproxylic insects.Wood-inhabiting fungi are accordingly better ableto colonize new suitable habitats, but still they areonly able to survive if the landscape as a whole offersa certain amount of suitable dead wood habitats. Abottleneck in dead wood abundance in Halland duringthe last centuries most likely has lead to the extinctionof some indicator species in Halland, e.g.
Stropharia hornemannii (stor kragskivling) is normally a fungus of coniferous forests, but in Halland itwas quite common on strongly decayed beech logs. It has not been recorded on beech wood in any of theNat-Man datasets, and seems to be a speciality for Halland in a beech forest perspective. Photo: JacobHeilmann-Clausen.
26
Saproxylic fungi
Ischnoderma resinosum which occur in several sitesin Skåne and Zealand as well as in the rainy beechforests of Slovenia. Within Halland the localizeddistribution of several indicator species seem to pointto landscapes with a higher degree of dead woodcontinuity than general in Halland. This most clearlyis the case in the Biskopstorp/Oskarström area, whichclearly should be the priority area for conservationof wood-inhabiting fungal communities on beechwood in Halland.
SummaryIn conclusion, it seems reasonable to assume that thecurrent distribution of indicator species withinHalland reflect the quality and abundance of currentdead wood habitats at local scale and the historicalcontinuity in dead wood abundance at landscapescale. The sites richest in indicators all have highconservation value, either due to a high localcontinuity in natural forest features (Mårås) and/ordue to their position in landscapes with spatio-temporal continuity in the presence of dead wood.
In the European perspective, the rather low numberof indicators recorded in the whole region partlyreflects the general, moderate level of dead wood
continuity at the landscape scale. Climatic con-straints, however, seem to play an important role forseveral indicators, meaning that the potential maxi-mum indicator score in Halland are lower than inother parts of Europe.
The high frequencies of Dentipellis fragilis, Hericiumcoralloides, Multiclavula mucida and Pluteusumbrosus in Halland suggest that the beech forestsof Halland in fact have a somewhat higher con-servation value than the raw indicator scores suggest.The presence of the rarely recorded polyporesSkeletocutis vulgaris and Tyromyces sp. points in thesame direction.
AcknowledgementsI wish to thank Örjan Fritz for introducing me to theHalland beech forests, for substantial practicalsupport throughout the project, for company inseveral of the extensive study sites and for setting upthe report. Erik Aude is thanked for collaborating inthe project and for company in the field. Finally,Morten Christensen is thanked for joining me in thestart up of the project, while Gro Gulden (Galerina)and Leif Örstadius (Psathyrella) are thanked foridentification of a few critical specimens.
Ascotremella faginea (dallerskål) on a beech log at Holkåsen. Photo: Örjan Fritz.
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Halland 2004
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Rolstad, J., Gjerde, I., Gundersen, V. S. & Sætersdal,M. 2002. Use of indicator species to assess for-est continuity: a critique. – Conservation Biology16: 253-257.
Ryvarden, L., Gilbertson, R.L. 1993-1994: EuropeanPolypores, Part 1 & 2. - Synopsis Fungorum 6,Fungiflora, Oslo.
Stoltze, M., Pihl, S., 1998. Rødliste 1997 over Plan-ter og Dyr i Danmark. - Miljø- og Energiminister-iet, Copenhagen.
Tortiè, M. 1998. An attempt to a list of indicator fungi(Aphyllophorales) for old forests of beech andfir in former Yugoslavia. - Folia CryptogamicaEstonica 33: 139-146.
Walleyn, R., Versteirt, V., Fayt, P., Branquart, E. 2004.Quelques champignons rares ou intéressants enhêtraie belge. - Miscellanea Mycologica 80-81:45-53.
Zar J.H. 1999. Biostatistical Analysis, Fourth edition.Prentice Hall International, Inc. New Jersey,USA.
Länsstyrelsen HallandEnheten för naturvård & miljöövervakning
Meddelande 2005:7ISSN 1101-1084
ISRN LSTY-N-M-2005/7-SE
Tryckt på Halmstads Tryckeri AB, Halmstad, 2005
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Halland 2004
Intensive study sites
Dömestorp is situated at the northern slopes ofHallandsåsen close to the E6. The studied stand isdistinctly influenced by former management, and ismostly homogenous and even-aged, consisting ofgenerally healthy trees. Dead wood is present inmoderate amounts, mostly due to break-down of sub-dominant or suppressed canopy trees, but also somelarge, dominant canopy trees have broken down inrecent years. Large, strongly decomposed logs arescarce. The biggest and most decayed trees aregenerally found in small spots along the northernborder of the area, often in small copses facingconiferous plantations or younger beech stands. Thesoils are generally richer than in other parts ofHalland, due to deposits of glacial tills containinglime-rich material (Fritz & Berlin 2002). Growthconditions for beech are accordingly comparablygood, and most trees are tall and rather fast grown.The oldest trees have been found to be about 200years old (Niklasson 2003), while the DBH of thebiggest study tree was 69 cm.
Dömestorp stands out as the intensive locality withthe lowest average species richness per tree and evenoverall species richness is low, considering thenumber of study trees (Fig. 3). The numbers of red-listed species and indicators are moderate (Table 3)pointing out the site to be of only moderate value forsaproxylic fungi. Notable records include Dentipellis
fragilis on two logs and Stypella subgelatinosa (syn.Protodontia subgelatinosa) on one. The latter ispresently only known from two other localities inSweden, and is new for Halland. Also the record ofthe little known Dacrymyces enatus is worthmentioning. Dömestorp is the top site for ecto-mycorhizal fungi in Halland, with occurrence ofseveral red-listed species (Fritz & Berlin 2002). Therichness of epiphytic lichens and bryophytes ismoderate, while the saproxylic insect fauna is poorlyknown. The rather limited value of Dömestorp forsaproxylic fungi most likely reflects former intensiveuse of the forests for lifestock grazing and/or timber/firewood production. The lack of well decayed logspoints to a break in the continuity of dead woodpresence, which may have been detrimental to someof the most demanding saproxylic species. In thecoming years the amounts of dead wood atDömestorp are likely to increase considerably. It willbe interesting to follow if saproxylic diversity willbe restored in the coming years as dead woodamounts increase.
The Biskopstorp areaThe landscape of Biskopstorp is situated 10-15 kmN of Halmstad and is the best preserved example ofa Mellanbygd forest landscape in Halland. Thenumber of woodland key habitats, especially old-grown deciduous forests, is exceptional and the num-ber of red-listed species unsurpassed in Halland (Fritz2004a). A 865 ha reserve is presently being designed
in the area. Four sub-localities inthe Biskopstorp area werestudied intensively in the project,while two sub-localities wereinvestigated shortly at thesummer-visit. All sub-localitiesare situated within a circle witha diameter of 2 km, and areconnected by more or lesscontinuous beech forests. Thismeans that exchange of fungal
Appendix 1. Description and notes on localities studied inthe project
A typical forest interior fromDömestorp, characterized byeven-aged, tall and mostlyhealthy beech trees. However,Lobaria pulmonaria and someother redlisted lichens grow onthe tree in the foreground.Photo: Örjan Fritz.
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Saproxylic fungi
spores is likely to be almost unhindered and contin-uous between the studied sublocalities. It is thereforeconsidered most likely that populations of mostsaproxylic fungi not strongly divided insubpopulation. In a European perspective it istherefore reasonable to treat all sub-localities in acollective sense. Doing this, the number of indicatorsadds up to seven, according to the European indicatorscheme (Christensen et al. 2005), placing Biskopstorpamong the most important areas for fungi inhabitingdead beech wood, known so far in Sweden (Table11). No other study localities in Halland hosted morethan four indicator species, though intensified studiesmay increase the number of indicators in several sites.Also for red-listed species Biskopstorp is outstanding,with records of 13 species associated with decayingbeech wood, including Ceraceomyces sulphurinus,which was found for the first time in Sweden since1940 (Table 3).
Holkåsen is the most intensively surveyed of the sub-localities at Biskopstorp, with 50 study trees. The
area consists of a rounded ridge covered by up to300 years old beech trees (Bengtsson 1999a). Thestand is fairly even-aged, with an average age closeto 260 years, and partly very open due to collapse ofold beech trees and removal of scattered Norwayspruce. Other parts are still closed forest, andespecially the northern, north exposed parts are wellprotected from desiccation from wind. The humuslayer is mostly shallow and growth conditions arefairly marginal except for a few spots with deepersoils at the bottom of the ridge. The DBH of thebiggest study tree was 70 cm, but most of the treeshad a DBH well below 60 cm.
Holkåsen stands out as the most species rich of allstudy sites (Fig. 3). Ten redlisted species wererecorded (Table 3). Most notable is the presence ofVolvariella bombycina, which was not found in otherlocalities during the project. Hericium coralloideswas recorded on five trees, while Multicalvulamucida was recorded on not less than 30% of thestudy trees. This means that Holkåsen may host the
Snags, partly decayed by Fomes fomentarius, are a very characteristic aspect of old-grown beech forestsin Halland, as here in Holkåsen. The upright position of snags causes a drier microclimate than in fallenlogs, and this seems to slow down the decay process. Several species seem to prefer standing snags, e.g.Camarops tubulina, Hericium coralloides, Phleogena faginea, while others typically sporulate in abun-dance after a standing snag has broken down, e.g. Nemania atropurpurea and Sistotrema sernanderi.Photo: Örjan Fritz.
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Halland 2004
biggest population of this species in the whole ofsouthern Scandinavia. Apart from saproxylic fungithree uncommon ectomycorrhizal fungi wererecorded viz. Cortinarius anthracinus, C. sanguineusand Lactarius fuliginosus. The locality is of out-standing value for saproxylic insects (Jansson 2004)and of high value for epiphytic lichens andbryophytes (Fritz 2001). The high biodiversity valueof Holkåsen does without doubt reflect the unusualhigh tree age, the high amounts of dead wood in allstages of decay, and the high variation in edaphicconditions.
N Kroksjön is situated just NE of Holkåsen whichare separated by a narrow stretch of natural bog. 25trees were studied. The area is very heterogeneous.The SW part is a thinned stand of well grown, rathertall beech which was managed until recently (1992).The central part consists of lower, more slowly growntrees on a low granite knoll, while the NE part is ona low-lying plateau. Parts of the latter area areunusually dark and closed for the area, while thecentral and SW part is more open due to thinning,poor growth conditions or, more locally, due tonatural breakdown of smaller tree groups. The humus
layer is well developed in most parts and the growthconditions for beech are generally more favourablethan in Holkåsen. Several trees with DBH exceeding60 cm are included in the study, with a maximum of92 cm.
N Kroksjön hosted a remarkable high number of red-listed species and was the only locality with morethan four indicators species following the Europeanindicators scheme (Table 3). Two of these, Inonotuscuticularis and Volvariella bombycina, were notrecorded in the present study, but are likely still tothrive at the locality. The very high value of NKroksjön for saproxylic fungi is somewhat surprisingconsidering the rather distinct signs of recentmanagement. However, the bigger size of study treesmay benefit some demanding heart-rot agents andthe vicinity to Holkåsen secures an unhinderedexchange of spores between the two localities. Thelocality is of rather high value for epiphytic lichensand bryophytes (Fritz 2001), while the fauna ofsaproxylic insects is unknown.
Kvinnsåsen is situated about 2 km NE of Holkåsenon a rather steep W to NW-facing slope. 15 trees
Small and very slow-grown beeches like these in N Kroksjön are characteristic for several of the oldgrown beech forests in Halland. They point to very poor soil conditions and/or an earlier history of coppi-cing and livestock grazing. Photo: Örjan Fritz.
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Saproxylic fungi
were studied. The stand is uneven-aged and in mostparts dense, closed and rather dark with a moistmicroclimate. The oldest trees are recorded to beabout 270 years old (Bengtsson 1999a). Growthconditions for beech are poor to average and the treesare mostly rather small and slow-grown. The DBHof the biggest study tree was 52 cm.
Six red-listed species were recorded at Kvinnsåsen(Table 3) of which Multiclavula mucida is the mostnoticeable. Other interesting records are Hohen-buehelia fluxilis and Omphalina epichysium whichwere found on a single log each. The locality had alow score of indicator species, and is at present, theleast valuable of the intensive sub-localities inBiskopstorp. This probably reflects the rather lowamounts of dead wood and the rather smalldimensions of most fallen trees.
Trälhultet is situated adjacent to Kvinnsåsen, onrather steep SE facing slopes. Ten trees were studied.The stand is distinctly influenced by formermanagement (up to 1992), and consists of large andtall, widely spaced trees. The oldest trees are recordedto be between 200 and 250 years old (Bengtsson1999). Dead wood amounts are moderate, but severallarge trees in different decay stages are present.Growth conditions are comparatively favourable,thanks to the warm, local microclimate and the ratherdeep developed soils. Several study trees had a DBHexceeding 60 cm, with a maximum of 78 cm.
Seven red-listed species were recorded, includingHericium coralloides, Lentinellus vulpinus and richoccurrences of Multiclavula mucida. Othernoticeable records include Lentinellus ursinus andOxyporus corticola. The latter species was notrecorded in any other localities during the project,while the two Lentinellus species were not found inany other sites in the Biskopstorp area. The high shareof red-listed species and indicators in Trälhultet isremarkable considering the small size of the area andthe moderate amounts of dead wood. The generallarge size of the trees may be one explanation forthis, but the favourable, warm microclimate may alsobenefit some specialised heart-rot agents. It seemslikely that more rare species may appear in Trälhultetas dead wood amounts increase in the coming years.
NÖ Kroksjön and S Ry were visited only brieflyduring the summer visit, and very few indicators orred-listed species were recorded. Most remarkableis the find of the strongly yellow corticoidbasidiomycete Ceraceomyces sulphurinus which isknown from very few sites in Scandinavia.
ÅkullaThe Åkulla area is situated about 15 km E of Varbergin a landscape rich in small lakes and steep ridges.The landscape hosts the northernmost concentrationof old-grown beech forests in Halland, and woodlandkey habitats are well represented (Fritz & Larsson2000). Two intensive localities were studied in thearea. They are situated about 2.5 km apart, separatedby farmland, coniferous plantations and swamps/lakes. It is therefore considered most likely that theyare functioning as discrete entities with only limitedexchange of fungal spores. For this reason they arenot treated in a collective sense with respect to theEuropean indicator scheme, as was done with thestudy sites in Biskopstorp.
Valaklitt is situated on rather steep, S-facing slopesfacing a small lake. The stand is rather heterogeneous.Large parts are dominated by collapsing, overmaturetrees and several very large sun-exposed canopy gapsare prominent. However, regeneration is welldeveloped, and other parts of the stand are dominatedby young or mature trees. As a whole the stand givesthe impression of very high naturalness with apronounced, though large-grained mosaic-structurewith well developed patch-dynamics as described e.g.from Suserup Skov in Denmark (Emborg et al 2000).The soils are partly deep developed, resulting inrather good growth conditions for beech. Severalstudy trees had a DBH exceeding 60 cm, with amaximum of 69 cm. 25 trees were studied.
Six red-listed species were recorded at Valaklitt(Table 3). Of these, Dentipellis fragilis, Camaropstubulina and Hercium coralloides were recorded onthree trees each. The latter was in one case found incompany with Inonotus obliquus which is an almostobligate associate in Danish beech forests (Heilmann-Clausen & Christensen 2000b). Hohenbueheliaauriscalpium, which is not previously published fromSweden, was found on two logs, while Meripilusgiganteus, which is very scattered in Halland, wasrecorded on one tree. With four indicators, followingthe European indicator scheme (Christensen et al.2005), Valaklitt is clearly among the most valuablesites for saproxylic fungi in Halland. It is most likelythat more indicators and red-listed species could beadded by inventories in the following years. The highvalue of the site reflects the long history of non-intervention and the high abundance of dead woodin all stages of decay. In addition the dominant SE-facing slopes may secure a rather warm microclimatebeneficial to some demanding species.
NV Äskemossen is very different from Valaklitt.
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Halland 2004
Most of the locality is situated on a low, undulatingplateau with small wet depressions and streams. Thestand is mostly dense and rather dark with only smallcanopy gaps. The amounts of dead wood aremoderate, and most dead trees are of fairly smalldimensions. Growth conditions for beech are poorto moderate and most trees are small and slow-grown.This probably reflect nutrient poor, partlywaterlogged soil conditions or the effects of formerheavy livestock grazing. The largest study tree had aDBH of 56 cm. 25 trees were studied.
Six red-listed species were recorded at the locality,most on single or very few trees. Most interesting isthe recording of Pluteus umbrosus which was notfound in Valaklitt. The corticoid Sistotrema radu-loides was found on a natural stump, associated withCamarops tubulina. The species is rare in all partsof Scandinavia, red-listed in Finland (Kotiranta &Niemelä 1996), and according to Hallingbäck &Aronsson (1998) not previously recorded in southSweden. The number of indicators in Äskemossen israther low and the value of the site for saproxylicfungi is at present only moderate. It is possible thatthis will change as dead wood amounts increase inthe future.
Extensive study sites
The Oskarström area
Getabäcken is situated about 10 km north ofOskarström in a densely forested landscape. Thelocality was investigated rather briefly and severallarge beech logs were not surveyed in detail.
The locality is dominated by rather closed and darkstands of tall beeches growing on gentle W-facingslopes. Locally, steep, exposed cliff-faces occur. Theforest structure gives an unusual natural impression,being uneven-aged with a well developed mosaic-structure. According to Niklasson (2003) the oldesttrees are about 285 years old. Growth conditions forbeech are favourable and most trees are unusuallytall and thick for Halland. Dead wood is present inmoderate amounts with a good balance betweendecay stages. A majority of the dead wood is presentas large logs. Uprooted trees are common, pointingto a deep developed soil layer and perhaps to Fomesfomentarius being less dominant than typical inHalland.
NV Äskemossen in the Åkulla area is a rather small, old grown beech stand dominated by slow-grownbeech trees of rather small dimensions. Photo: Örjan Fritz.
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Saproxylic fungi
Four red-listed species were recorded, including therichest occurrences of Dentipellis fragilis (sevenlogs) and Camarops tubulina (three logs) met within the project. The possibly undescribed polyporeTyromyces sp. was seen on four logs. Although thenumber of recorded indicators is rather low the richoccurrence of the above mentioned species points tovery high mycological values. The visit at Getbäckenwas rather short and it is highly probable that moredetailed investigations would lead to records of morerare species. The locality is very rich in red-listedsaproxylic insects and second in Halland only toHolkåsen in Biskopstorp in conservation value in thisrespect (Jansson 2004).
Myskebackarna is situated 5 km WSW of Geta-bäcken, closer to the arable farmlands along Kattegat.The locality was investigated rather throughoutly butsome of the smaller dead trees were not surveyed.The locality is dominated by rather low, often multi-stemmed beech trees on three rocky hills. Many treesshow distinct signs of former coppicing and indivi-dual stems are generally small. The amounts of deadwood are rather low to moderate, with a dominanceof recently dead wood. The oldest trees are dated to
be 240 years old, but most are almost 100 yearsyounger (Niklasson 2003).
Four redlisted species were recorded at the locality(Table 3). Except for Hypoxylon cohaerens all wererecorded only once, within a small area (“provplats4” in Niklasson 2003, p. 33). Hericium coralloidesand Ossicaulis lignatilis were in fact found on thesame log, meaning that the three most interesting red-listed species were recorded on only two logs! In thesame area Inonotus obliquus was recorded on an oldbeech snag while Mycena crocata was found onbeech litter. Inonotus obliquus is generally rare onbeech and mainly confined to beech forests with highdead wood continuity (Heilmann-Clausen & Christ-ensen 2000b). Mycena crocata is quite seldom in theregion, and mainly restricted to beech forests on richsoils. The highly localised co-occurrence of severaldemanding species is striking. The area is S-facingand stands out from the rest of the locality by theoccurrence of some large standing beech snags, anda relative richness of large, living beech trees. It isunclear if the local mycological values reflect aspecial local forest history or special growthconditions on the steep south-facing slopes. Jansson
Except for the rocky outcrops Getabäcken almost gives the impression of a Danish beech forest on ratherrich soil dominated by tall and large trees. Photo: Örjan Fritz.
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Halland 2004
(2004) found the locality in general to be the leastvaluable site for saproxylic insects among 15 beechforests inventoried in Halland. This corresponds wellwith the overall impression of the value of the sitefor saproxylic fungi, but it is unclear if any insecttraps were placed in the small area where theinteresting fungi were recorded.
Frodeparken is a small beech forest reserve on flat,deep developed soils midway between Biskopstorpand Myskebackarne. The site was visited ratherbriefly, but almost all dead trees were inventoried.The growth conditions for beech are favourable andthe locality is characterized by unusually tall and bigbeech trees. The stand is about 200 years old
(Niklasson 2003), fairlyeven-aged and homogene-ous. Dead wood amounts aremoderate to high and severallarge gaps occur in thecanopy.
Despite the high dead woodvolumes only one red-listedspecies were recorded at thelocality and even the numberof indicators is remarkablylow (Table 3). The area isrich in old stone-heapspointing to former farmingand a break in forest con-tinuity. This may explain thelack of rare fungal species,though it should be re-marked that the richness ofred-listed saproxylic insectsis rather high (Jansson2004).
Skrockeberg consist ofmainly E and SE-facing,rather steep slopes situatedin the densely forestedlandscapes north of Nissa-ström. The site was investi-gated rather briefly, but mostdead trees were surveyed.The forest is unusually richin large pines but alsospruces occur scattered. Thetrees are generally between
150 and 200 years old. Some of the pines showdistinct signs of forest fires in the 19th century,probably of anthropogenic origin (Niklasson 2003).Niklasson (2003) interpret the stand to be generallya secondary natural forest, unmanaged since livestockgrazing was abandoned 150-200 years ago. Five red-listed saproxylic fungi are known from Skrockeberg,including Camarops tubulina, Hericium coralloidesand Multiclavula mucida which were all found inthe present investigation, the latter on at least fourtrees. Dentipellis fragilis was not found in the presentinvestigation, but have been seen in recent years.Other interesting records include the possiblyundescribed Tyromyces sp. on at least two beech logs.Skrockeberg has a rich fauna of saproxylic insects,including several red-listed species (Jansson 2004),and is even rich in epiphytic lichens (Niklasson2003).
The beech trees in Halland are generally of rathersmall dimensions. More scattered, like in Nissa-ström, much larger trees occur, due to more favour-able soil conditions. Photo: Örjan Fritz.
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Saproxylic fungi
Nissaström is located along the banks of the riverNissan, 5 km north of Oskarström. The locality wasvisited rather briefly. It is a very heterogeneouslocality with several small areas with old beech forest,intermixed with oak forest and mixed deciduousforest including alder, lime and aspen. Parts of thelocality show distinct signs of former human use,but the steepest cliffs along Nissan has probablyalways been forested. The amounts of dead woodare rather low, but several unusually large beech logsoccur. The soils are partly very rich for Halland, withoccurrence of several ectomycorhizal fungi with avery restricted distribution in Halland, e.g.Cortinarius citrinus, Craterellus cornucopiae andHygrophorus nemoreus.
Three red-listed fungi were found on dead beechwood in Nissaström (Table 3), including Pezizasaniosa, which has not previously been recorded fromHalland. The species normally grows on rich soil,but is known from dead beech wood also in Denmark.Two other red-listed saproxylic fungi are known fromNissaström, i.e. Dentipellis fragilis and Clavicoronapyxidata but occurring on other hosts (lime? and
aspen, respectively). Only the former was recordedin the present investigation. Also the possiblyundescribed polypore Tyromyces sp. was found (onone log), while Galerina pallida was recorded as newto Sweden. Despite the distinct human impacts,Nissaström is a locality of considerable importancefor saproxylic fungi. This probably reflect the unusualsize of the trees at the site, the rich soil conditions,the local warm microclimate on the steep cliff-facesand probably a high, local dead-wood continuity onthe steep cliffs.
Råmebo is an old beech forest on rather flat terrain10 km SE of Oskarström. Half of the stand wasthinned and cultivated for regeneration 10-20 yearsago while another half part has been untouched forseveral decades. The canopy layer is fairly even-aged,consisting of tall, 250-280 years old trees (Niklasson2003). Dead wood amounts are moderate, but mostdead wood is present as large logs and snags. Mostof the fallen logs were investigated, except for thethinned parts where vigorous beech regenerationmakes most logs virtually inaccessible.
Unlike the situation in many other beech forest reserves in Halland, spruce is allowed to grow freely inRågetaåsen. As a result the forest is rather dark with several very large spruces and rich reproduction ofspruce offering serious competition for the beech saplings Photo: Örjan Fritz.
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Halland 2004
Four red-listed, saproxylic fungi are known from thelocality, including Dentipellis fragilis, Hericiumcoralloides and Lentinellus vulpinus (Table 3). Thelatter was found on three trees in the presentinvestigation, while Hericium coralloides is onlyknown from earlier years. Other noteworthy recordson beech include Inonotus obliquus, Lentinellus ur-sinus and Pluteus pellitus. The indicator scores areamong the highest for extensive localities in theproject (Table 3) and the locality is among the mostvaluable in Halland also for saproxylic insects,epiphytic lichens and bryophytes (Fritz 2001, Jansson2004). This points to high, unbroken forest continuity,even in the presence of dead wood and veteran trees.
Rågetaåsen is situated 10 km NE of Oskarström onsteep, rocky ESE-facing slopes. The locality ischaracterized by several small streams and bogs andthe presence of several very large spruces. Parts ofthe stand are probably in a successional change fromopen beech-forest with scattered spruce to a mixedconiferous stand, where beech is likely to play onlya subordinate role.
Niklasson (2003) found the oldest trees to be only130 years old, but this probably exclude the oldestgeneration of beech trees which occur as scattered,mostly dead or dying veterans. The stand wasprobably open grassland with only scattered treesuntil the mid part of the 19th century (Niklasson2003). Dead wood amounts are moderate, with a goodbalance between dead wood in various decay stages.The locality was visited rather briefly, but most ofthe oldest beech trees were investigated carefully.
Four red-listed species were recorded, with Gloeo-hypochnicium analogum and Dentipellis fragilis asthe most noteworthy species. Even the possiblyundescribed Tyromyces sp. was recorded on the baseof a rotten beech snag. Gloeohypochnicium analogumis known from very few records in Sweden. Thespecies has a southern distribution in Europe and itspresence in Rågetaåsen may be of relict character,pointing to warmer, more light-open conditions inthe past. It is doubtful whether any of the abovementioned species will be able to persist as the standsgets cooler and darker due to the vigorous growth ofspruce and the dieback of the old beech generation.
Mårås is clearly the most valuable inland locality for wood-inhabiting fungi in Halland. Parts of the standgive a very natural impression, with a well developed mosaic structure and a good balance betweenregeneration and breakdown of old trees. Photo: Örjan Fritz.
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Saproxylic fungi
Svarta Klippan is situated 7 km E of Råmebo facingthe small river Fylleån. The studied area is situatedon very steep W facing rocky slopes, and consists ofmixed deciduous stands with oak and beech beingdominant. Generally the growth conditions for beechare rather poor, but locally, close to sheltering rocks,larger trees occur. The oldest beech-trees are datedto be about 300 years old (Niklasson 2003). The steeppart of the locality with the oldest trees wasinvestigated throughoutly.
Three red-listed saproxylic fungi are known SvartaKlippan (Table 3). Most interesting is the presenceof Inonotus cuticularis and Gloeohypochnicium ana-logum. Both have a southern distribution inScandinavia and their presence points to a local,warm microclimate on the steep cliff-faces. Forsaproxylic insects Svarta Klippan is only of ratherlimited value (Jansson 2004) which may indicate abreak in dead wood continuity. Dead wood amountsare still rather sparse and it is interesting if more rarespecies will occur at the locality when more of theold trees collapse in the future.
Inland area (municipality of Hylte)
Mårås is situated right north of Hyltebruk and is thebiggest and by far the most natural of the beech forestreserves in the eastern part of the county of Halland(belonging to the landscape of Småland). The localityincludes 6-7 ha of very natural looking beech forestwith a well-developed mosaic structure and a goodbalance between tree generations. The most naturalarea is situated on a rather low hill and has allexpositions. The oldest dated beech in Sweden wasfound in the area in 2001. It was at that time dated tobe at least 400 years old, but died in 2002 (Niklasson& Fritz 2003). Several other trees in the stand areknown to be more than 300 years old (Niklasson2003). Growth conditions are generally fairly goodand many trees are tall and rather thick. Dead woodamounts are high and show a very good balancebetween decay stages. The locality was investigatedrather throughoutly, but only the most interestinglooking logs were inventoried carefully.
With seven red-listed species Mårås is clearly thetop scorer among the extensive localities. Dentipellisfragilis, Hercium coralloides and Pluteus umbrosus
The inland locality Skubbhult host rich communities of redlisted lichens and saproxylic insects, but onlyfew interesting wood-decay fungi were recorded. Photo: Örjan Fritz.
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Halland 2004
were each recorded on two or three trees, whileCamarops tubulina and Multiclavula mucida wererecorded on a single tree each. The latter was foundin very sparse amounts on a dead log adjacent to thesouthern border of the forest facing open land. Thespecies has not previously been recorded at Måråsand is probably in the phase of colonization. Theoccurrence of wood decaying agarics was remarkablyrich, with high frequencies of Omphalina epichyisumand several Pluteus species. Mårås has a very richflora of epiphytic lichens (Niklasson & Fritz 2003),while the fauna of saproxylic insects is rather poorlydeveloped (Jansson 2004).
The discrepancy between the richness of saproxylicinsects and fungi in Mårås is intriguing and hard toexplain. It is worth noticing that the interesting wooddecay fungi were found very scattered across thewhole locality, while most of the other top sites inHalland were characterized by an often strikinglylocalized co-occurrence of several rare species. Inother words, it is possible that the more natural foreststructure in Mårås means that the most demandingspecies occur more scattered, because their requiredsubstrates occur scattered. This contrasts withlocalities characterized by more even-agedgenerations of very old trees offering local hotspotsof substrate abundance. If the differences in foreststructure has this effect, it is likely that the ratherlow number of insect traps in Mårås has been able tocatch up with only a limited part of the total insectsdiversity at the site. The fungal inventory, in contrast,covered the whole locality and most likely recordeda very high fraction of the total diversity.Alternatively, it is possible that saproxylic insectsand fungi differ in important habitat requirementsand therefore have responded differently to breaksin forest continuity and/or have variable affinity tothe present habitat conditions.
Skubbhult is situated next to the lake Mellan-Färgenright north of Femsjö parish; the birth place of EliasFries. Three adjacent beech stands were studied. Mosttime was used in the central part of the area(corresponding to “kärnområdet” in Fritz (2001)),while two areas closer to the lake were inventoriedless carefully. The central stand is in almost levelterrain and gives a fairly even-aged impression.Niklasson (2003) found it to be about 170 years old,with very few older trees. The amounts of dead woodare moderate and newly dead wood predominate.
Only two red-listed species were recorded inSkubbhult, with Phleogena faginea being the mostinteresting. Even the numbers of recorded indicators
are low and as a whole Skubbhult seem to of limitedimportance for saproxylic fungi. In contrast thelocality is rich in red-listed epiphytic lichens andbryophytes and saproxylic insects (Fritz 2001,Jansson 2004).
Ödegärdet is the eastern-most of the study sites inthe municipality of Hylte. The locality is quiteheterogeneous but most of the study area consists oflow-grown, fairly even-aged beech forest. Dead woodamounts are generally low to moderate, and mostdead wood are of rather recent date. The area showdistinct signs of former human use and is an importantarchaeological locality. At least parts of the site havebeen used for farming at various times during thelast 2000 years. Most of the old beech trees arebetween 200 and 250 years old, while a few fallenveteran oaks are considerably older (Niklasson 2003).
No red-listed fungi were recorded in Ödegärdet andeven the number of indicators is strikingly low (Table3). The locality was visited rather briefly under poorlight conditions and some species may have beenoverlooked. Nevertheless it seems clear that the siteis of limited value for saproxylic fungi, whichprobably reflect breaks in dead wood continuity inthe past. Some of the slopes facing the small roadeast of the locality were found to host interestingectomycorrhizal fungi including Cortinariusvariegates and Tricholoma portentosum which areonly rarely found in pure deciduous forest.
Southern Halland
Blåalt is a rather big, but fairly isolated forest reserveSW of Mästocka in the southern part of Halland. Thearea is very heterogeneous and includes mixed deci-duous stands with oak, beech and pine as well aspure beech forests. The locality was visited both atthe summer and the mid autumn visit. At the latteroccasion only parts with large amounts of dead beechwood were inventoried, i.e. the western-most part ofthe reserve S of the parking place and the areas alonga small ravine in the central part of the reserve (SEof “provplats” 3 in Niklasson (2003)). In both areasmany beeches show distinct signs of former coppi-cing. Some are multi-stemmed, while others havebasal scars and cavities pointing to old wounds. Theoldest beech stems was dated by Niklasson (2003)to be about 200 years old. The stands are still gene-rally closed and dead wood amounts are moderate.
Despite much time used in the field only few red-listed fungi were recorded at Blåalt (Table 3). More
40
Saproxylic fungi
noteworthy are the records of Coprinus extinctoriusand Entoloma placidum. Both are known from veryfew records in Sweden, but are normally not asso-ciated with beech forests of high conservation valuein Denmark (pers. obs.). The number of recorded indi-cator species is low and Blåalt has, as a whole, ratherlow to moderate value for saproxylic fungi at present.Most striking is the lack of several species associatedwith old fallen logs and snags, i.e. Camarops tubuli-na, Dentipellis fragilis, Hericium coralloides, Ino-notus cuticularis and Tyromyces sp. The locality isamong the most valuable sites for epiphytic lichensand bryophytes in Halland (Fritz 2001), while theimportance for saproxylic insects is relatively low(Jansson 2004).
Vallåsen is situated at the central part ofHallandsåsen, 10 km SE of the intensive study siteDömestorp. Vallaåsen is a fairly even-aged beechstand in undulating terrain. Most of the surroundingareas are covered by coniferous plantations and thestand is fairly isolated from other old beech stands.
The amounts of dead wood are high due to increasingbreak down of old canopy trees. Several large canopygaps occur and many logs are sun-exposed. The standis generally wide-spaced and most trees are tall andthick-stemmed. The oldest trees are about 200 yearsold (Niklasson 2003). The locality was visited bothat the summer and the mid autumn visit.
Four red-listed species were recorded, includingPluteus umbrosus, Hericium coralloides (two trees)and Lentaria epichnoa (three trees). The latter specieswas not recorded in other localities in the projectand is new to Halland. Interesting is also theabundancy of Peziza micropus which was notrecorded in the nearby intensive locality, Dömestorp.Also several Pluteus species were much moreabundant in Vallåsen than in Dömestorp. Vallåsenhas high value for saproxylic fungi but the small sizeand isolated position may in the long run causeproblems for the local survival of the presentpopulations of the most demanding species.
Several of the old-grown beech forests in Halland are marked by former management. This is also thecase in Vallåsen, which is characterized by a fairly even-aged, widely spaced generation of beech trees.The stand is now collapsing and a decrease in dead wood amounts are foreseeable within the cominggenerations. The massive front of conifers in the background indicates another problem. Many old grownbeech stands are isolated and surrounded by other habitat types which may hinder effective spore-disper-sal between suitable habitats rich in dead wood. Photo: Örjan Fritz.
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Halland 2004
Phleogena faginea (VU), pulverklubba, is a veryunusual heterobasidiomycete forming groups of tinyclub-shaped sporocarps, reminding of myxomycetefructifications. In cases of doubt a strong smell ofcurry may help in the identification. It is often foundon hard and dry wood, e.g. on rather fresh snags orunderneath recently broken logs. The species fruitslate in the season and was found in all intensivelocalities except Kvinnsåsen. Most likely it occurseven in many of the extensive localities, which wasvisited only in mid-autumn. The species wasespecially abundant in Holkåsen, where it was foundon 20 % of the studied trees. The species is consider-ed very rare in most parts of Europe, but is most likelywidely overlooked due to the inconspicuous sporo-carps, formed late in the year. The species is not pre-viously recorded from Halland, and according torecent information from Anders Dahlberg at “Art-Databanken” the species was known from only tenlocalities in Sweden prior to the present investigation.It was listed as an indicator of habitat quality inDanish beech forests by Heilmann-Clausen &Christensen (2000b), but omitted from the Europeanindicator system (Christensen et al. 2005), due to the
rather inconspicuous sporocarps and uncertaintyregarding ecological preferences.In Britain more than100 records are known, some made on oak saplings(Martyn Ainsworth pers. comm.)
Stypella subgelatinosa (syn. Protodontia subgelatin-osa)(NT), lövgråtagging, is a heterbasidiod, corticoidfungus forming masses of tiny greyish spinesunderneath dead wood. It was found as new toHalland in Dömestorp. The species is red-listed alsoin Denmark (Stoltze & Pihl 1998) and Norway(Bendiksen et al. 1997). It inhabits strongly decayeddeciduous wood and probably has a preference forricher soils. It is found in all parts of Scandinavianorth to Torne Lappmark. The sporocarps are veryinconspicuous and the true rarity of the species isdebatable.
Ceraceomyces sulphurinus (DD), svavelskinn, is acorticoid fungus forming conspicuous stronglyyellow sporocarps with numerous mycelial cords.The species was during the project found for thesecond time in Sweden. The first record dates backto 1940, when the species was found close to Femsjö.
Appendix 2. Notes on red-listed and other interesting speciesrecorded in the project.
Dentipellis fragilis (skinntagging) occurred in abundance on fallen beech snags in Getabäcken.Photo: Örjan Fritz.
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Saproxylic fungi
Hericium coralloides (koralltaggsvamp) is unmistakeable due to its delicate branched, coral-shapedsporocarps. Nissaström/Getabäcken. Photo: Örjan Fritz.
Outside Sweden the species is in Scandinavia knownfrom a few localities in Finland. The new record wasmade in NÖ Kroksjön at Biskopstorp underneathstrongly decayed wood of beech. In the field thespecies were taken for Phlabiella vaga which formsimilarly coloured sporocarps. The species might bequite distributed in Halland as similar lookingsporocarps were seen at the summer visit also inBlåalt and Trälhultet, but not collected. At the mid-autumn visit the species was not found again, andthe species may have a rather short period ofsporulation.
Gloeohypochnicium analogum (syn. Hypochniciumanalogum) (DD), lundkrämskinn is a corticoid fungusforming rather thick, pinkish cream sporocarps witha powerful fruity smell. It was recorded as new toHalland with finds in Rågetaåsen and Svarta Klippan.The species has a southern distribution in Scandi-navia with very scattered records from Denmark,Skåne, Blekinge and the Mälaren-area in Sweden. Itwas listed as an indicator of habitat quality in Danishbeech forests by Heilmann-Clausen & Christensen(2000b), but omitted from the European indicatorsystem (Christensen et al. 2005), due to the ratherinconspicuous sporocarps. In contrast, Ainsworth
(2004) still list it among the indicators for importantsites for saprotrophic fungi on beech in the UK.
Sistotrema raduloides is a corticoid fungus formingmasses of small, whitish to pale brownish spines ona slightly paler subiculum. It was found as new toHalland at Äskemossen on a rotten beech snag. Thespecies is red-listed in Finland (Kotiranta & Niemelä1996), and according to Hallingbäck & Aronsson(1998) not previously recorded in south Sweden.
Dentipellis fragilis (NT), skinntagging, grows ondeciduous wood and is in Scandinavia most commonon beech and aspen. It was recorded from eight ofthe study sites but occur rather widespread in thecounty of Halland (Fig. 14). The largest sporulatingpopulation was encountered in Getabäcken, wheresporocarps were seen on at least seven trees. It wasnot found in the Biskopstorp sites, and is notpreviously recorded in the area. This is surprisingconsidering the general high fungal diversity of thearea. Dentipellis fragilis is red-listed also in Denmark(Stoltze & Pihl 1998), Norway (Bendiksen et al.1997) and Finland (Kotiranta & Niemelä 1996), listedas an indicator of biologically valuable old growthforests in both Estonia (Parmasto & Parmasto 1997)
43
Halland 2004
and the former Yugoslavia (Tortiè 1998), andincluded among the European beech forest indicatorsby Christensen et al. (2005). The species has aneastern distribution in Europe and is not known fromthe UK. In Halland it was mainly found on standinghigh snags or recently downed logs/snags, which hadbeen standing dead for several years. Only in onecase it was associated with a log strongly infectedwith Fomes fomentarius. Details on the ecologicalrole of the species and its infection strategy are stillunknown.
Hericium coralloides (NT), koralltaggsvamp, is awell known and beautiful decayer of deciduous wood.In Sweden it occurs mainly on beech, birch and aspen(Hallingbäck & Aronsson 1998). It was recorded from12 study sites and is a fairly widespread species inHalland (Fig. 14). The richest occurrence was foundin Holkåsen. The species is red-listed in mostEuropean countries and considered as an importantindicator of old natural forests in Denmark, Estonia,Poland, UK and the former Yugoslavia (Parmasto &Parmasto 1997; Tortiè 1998; Heilmann-Clausen &Christensen 2000b; £uszyñski 2003; Ainsworth2004). It is even included among the European beech
forest indicators by Christensen et al. (2005). Allrecords were made on standing or recently downedbeech snags which had been standing dead for someyears. Hericium coralloides seem to have quite strictassociations with other wood decay fungi (Niemeläet al. 1995). In Halland it was in all cases found inassociation with Fomes fomentarius, while Inonotusobliquus and less often I. cuticularis are the mostcommon associated species in Denmark (Heilmann-Clausen & Christensen 2000a).
Lentinellus vulpinus (NT), rynkmussling is ratherclosely related with Hericium coralloides and allies.Its sporocarps are however of the agaric type, butrather tough and with irregular, serrated gills. Thecap is pale tan to pinkish brown with a distinctlyribbed cap-margin. It grows on various deciduoushosts. The species has been much confused with theclosely related L. ursinus, which differs in itsdistinctly brown-tufted pileus. The species was foundas new to Halland in Trälhultet and Råmebo. Thespecies is red-listed also in Denmark (Stoltze & Pihl1998) and Norway (Bendiksen et al. 1997) andincluded among the European beech forest indicatorsby Christensen et al. (2005). The record in Trälhultet
Lentinellus ursinus (björnmussling), showing the characteristic contrast between the dark felty centralpart of caps, and the paler and almost smooth margins. Råmebo. Photo: Örjan Fritz.
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Saproxylic fungi
were made on a still living but partly rotted beechtree while it was found on a standing, dead snag inRåmebo. It seems probable that the species formsheart-rot in living trees.
Lentinellus ursinus, björnmussling, has been muchconfused with the above species and L. castoreus. InSweden it has mostly been treated under the lattername (e.g. Hallingbäck & Aronsson 1998). For thisreason the true distribution of the species in Swedenis unknown. In the current circumscriptionLentinellus ursinus is defined by being distinctlybrown hairy to furry at the central part of the cap,strongly contrasting to the paler and smoother cap-margin. The species was recorded as new to Hallandon the same two localities as the above species, andone might wonder if they are really different at thespecies level. It is included among the Europeanbeech forest indicators by Christensen et al. (2005)and used also in regional indicator systems inBelgium and the UK (Ainsworth 2004; Walleyn etal. 2004). In Halland it was found on standing orfallen beech snags which had been standing dead forseveral years. In several cases it was occurring inlarge amounts. The same ecology seems to be typicalin Denmark and Fountainebleau in France, where thespecies is quite distributed (pers. obs). The speciesmay be a heart-rot agent of old, over-mature trees or
be dependent on one or more specific predecessor asHericium coralloides.
Ceriporia excelsa (NT), rosenporing, is a resupinatepolypore forming fragile, often widely effused,pinkish to violaceous or orange sporocarps. Thespecies is difficult to separate from the related C.viridians, which differ mainly by its duller and oftengreenish colours. It is possible that the two speciesare conspecific or belongs to a still unresolved speciescomplex. It grows on decaying deciduous wood andwas recorded from eight study sites in the project. InDenmark the species is rather common and notstrictly associated with valuable natural forests. It isnot included in the beech forest indicator schemesby Heilmann-Clausen & Christensen (2000b),Ainsworth (2004), Christensen et al. (2005), thoughit is regarded as a indicator of habitat quality inSweden (Hallingbäck & Aronsson 1998). In Hallandthe species was typically found on dead beech logsor branches in medium decay stages. It was notparticularly common in any of the study sites.
Inonotus cuticularis (VU), skillerticka, formsaggregated clusters of rusty brown sporocarps onliving and recently dead deciduous trees, particularlybeech. It was not seen fertile during the project, butold sporocarps were seen on two living trees; one in
The mysterious Tyromyces sp. photographed on a large beech log in Nissaström. Future work is plannedin collaboration with Professor Leif Ryvarden, Oslo, to sort out the status of the species. Photo: JacobHeilmann-Clausen.
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Halland 2004
N. Kroksjön and one in Svarta Klippan. There is littledoubt that the species still thrive in both trees, butprobably sporulating was not initiated in 2004,because of the rather cool and rainy summer. Thespecies has a southern distribution in Scandinaviaand is known only from one recent record in Norway(Bendiksen et al. 1997), while it is rather distributedin Denmark. In Halland it is known from fourlocalities in total (Fig. 13). It is included among theEuropean beech forest indicators by Christensen etal. (2005) and its presence in the two sites probablypoints to an unbroken continuity in the presence ofsuitable veteran trees and a favourable, warmmicroclimate at the local scale.
Inonotus obliquus, sprängticka, is very common onbirch in most parts of Scandinavia where it formsblack, asexual, irregular conks on living trees. Theporoid, resupinate sporocarps are formed lesscommonly on recently dead, fallen or still standingtrunks. On beech the asexual, black conks are neverseen and it is possible that the species growing onbeech is a distinct, undescribed taxa. In HallandInonotus obliquus was recorded on beech in threesites, in all cases on sun-exposed, standing snags.This ecological preference is very distinct for thebeech form in Denmark too (pers. obs) while thetypical birch form grows in all kind of expositions.The form on beech was included among the indicatorsof habitat quality in Danish beech forests byHeilmann-Clausen & Christensen (2000b) butomitted from the European system (Christensen etal. 2005) due to the unclear taxonomical status. Itwould be interesting to do mating and/or DNA studieswith both forms to resolve if they are distinct taxa ornot.
Skeletocutis vulgaris, sydlig gräddporing, wasrecently separated from the probable sib species S.lenis, gräddporing (Niemelä & Dai 1997). Both formwhite to cream-coloured, thin and strictly resupinate,poroid sporocarps on dead deciduous and coniferouswood. The ecology and distribution of Skeletocutisvulgaris is still poorly known, but the species seemsto be quite distributed in Halland, with records from10 study sites. It was mainly found on rather bulkysubstrates, both snags and lying logs. A preferencefor rather fresh, undecayed wood, as mentioned byNiemelä & Dai (1997), was not evident, as the speciesin several cases was found on very rotten wood. It isunclear if the species has a preference for sites ofhigh natural value, but it is noteworthy that the verysimilar S. lenis is included in the red-lists in bothSweden and Finland (Kotiranta & Niemelä 1996;Gärdenfors 2000). Skeletocutis vulgaris was not
recorded at all in the Nat-Man project (Ódor et al.2004a), and may be a Halland speciality in a Euro-pean beech forest perspective.
Tyromyces sp. Already before the beginning of thefield-work in Halland, Örjan Fritz mentioned theexistence of a strange, yellowish, unnamed polyporein some of the old beech forests in Halland. At themid-autumn visit I had the chance to study the speciesmyself on a total of seven large beech trees in fourdifferent localities, but could not suggest a plausiblename. Since then several Danish mycologists hasbeen consulted and dried material has been checkedby Professor Leif Ryvarden in Oslo. The preliminaryconclusion is that the species is undescribed or atleast not well known in Europe. Macroscopically thespecies has some resemblance to Tyromyces kmetiibut the spores are narrower than in this species. AlsoCeriporiopsis gilvescens is rather similar, but differsin the strictly resupinate sporocarps, a narrower pore-layer and pinkish colours. It is planned to assess thetaxonomic position of the species by DNA sequen-cing as basis for an eventual scientific description ofthe species as a new taxa.
The species seem to be restricted to the wettest partof study area, where it is known from five sites intotal (Fig. 13). It would be highly relevant toinvestigate further potential sites for the species.Known sites deserve the biggest possible conser-vation attention, as the species may be very rare atthe global scale.
Lentaria epichnoa (NT), vit vedfingersvamp, formstiny coral-shaped sporocarps on rotten deciduouswood. The species was recorded on three logs inVallåsen and is new to Halland. It is generally morecommon in central Sweden and is considered as agood indicator of habitat quality in aspen stands(Nitare 2000). In Denmark most of the few recordsare from heaps of sawdust, but it is also found ondecaying beech logs in two localities, both of highconservation value.
Multiclavula mucida (syn. Lentaria mucida) (VU),vedlavklubba, is a lichenized basidiomycete formingdense groups of small whitish club-shaped sporocarpson rotten wood, which is coloured green by the algalphotobiont. It was recorded in eight study sites andwas especially abundant at Holkåsen. The populationin Halland is restricted to the Biskopstorp area and afew sites north and east hereof (Fig. 13). The speciesis more distributed in central Sweden (Nitare 2000),with only very scattered records south of LakeVänern. The species most likely depends on a stable,
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Saproxylic fungi
Figure 13. Distribution maps of Tyromyces sp., Multicalvula mucida, Inonotus cuticularis and Pluteusumbrosus in Halland, based on this and previous investigations. All have a very restricted distribution inHalland. The two first mentioned are recorded mainly in the Oskarström area, while the two latter areeach recorded from about a handful of localities scattered over the county.
Tyromyces sp. Multiclavula mucida
Inonotus cuticularis Pluteus umbrosus
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Halland 2004
Figure 14. Distribution maps of Hericium coralloides, Dentipellis fragilis, Camarops tubulina and Hypo-xylon cohaerens in Halland, based on this and previous investigations. All occur fairly widespread in thecounty.
Hericium coralloides Dentipellis fragilis
Camarops tubulina Hypoxylon cohaerens
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Saproxylic fungi
The delicate sporocarps of Lentaria epichnoa (vit vedfingersvamp) was found on several stronglydecayed beech logs in Vallåsen. Photo: Jacob Heilmann-Clausen.
Multiclavula mucida (vedlavklubba) growing on its green mat of algae on strongly decayed beech woodin Holkåsen. Photo: Jacob Heilmann-Clausen.
humid forest climate, and the rich occurrence incentral Halland probably benefit from the highprecipitation in the region. It is not known fromDenmark and was in the Nat-Man project only
recorded from Slovenia, where both the study sitesare in high precipitation areas. According to Holec(2003) it is restricted to forests minimally influencedby human activities in the Böhmerwald, the Czech
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Halland 2004
Republic, and should probably be regarded as aindicator of forests of high conservation value assuggested by Nitare (2000)
Bolbitius reticulatus (NT), hinnskivling, is a smallagaric reminding of a Pluteus but differing by theochre gills and the sticky pileus. It was found in sixstudy sites, in all cases on strongly decayed, mostlylarge beech logs. It is found north to northern centralSweden and seems to have a preference for richersoils (Hallingbäck & Aronsson 1998). It might be agood indicator of habitat quality in the region, thoughit is not included in any of the beech indicatorschemes used in the rest of Europe.
Coprinus spelaiophilus (syn. Coprinus extinc-toriusss. auct) is a rather large but thin-fleshed Coprinus.Macroscopically it is reminding of C. domesticus,but it differs among other things by the structure ofthe veil and the more pure white cap. It is typicallyassociated with hollows in living trees (Rald 1989)and the record in Blåalt was made in exactly thishabitat. The species is poorly known in Sweden, buta single record from Skåne is listed in Hansson &Hägg (2000). It seems to be a rare species in Europe(Rald 1989) and is included in the Danish red-list(Stoltze & Pihl 1998). The indicator value of thespecies is unknown but probably limited as it oftengrows in solitary trees in cities and parks (Rald 1989).
Galerina pallida is a very unusual Galerina charac-terized by striking pale colours and hyaline, dex-trinoid spores. For this reason it was first describedin its own genus as Velomycena pallida. InScandinavia it is known only from a few finds inNorway (Gro Gulden pers. comm.) and is hererecorded as new to Sweden. It was found on rottenbeech wood in Nissaström, and was in the fieldmistaken for an albino form of G. marginata. In theNat-Man project it was even recorded on beech woodin the Slovenian virgin forest site Rajhenavski Rog(Piltaver et al. 2002). The present collection wasdetermined by Gro Gulden, Oslo.
Galerina pseudomniophila & G. vexans are twobryophyte associated Galerina’s which are herepublished as new to Sweden, thanks to determinationwork by Gro Gulden. Galerina pseudomniophilaforms small yellow brown sporocarps reminding ofG. mniophila, but characterized by a slightly differentmicroscopy. It was recorded from Vallåsen and N.Kroksjön. Galerina vexans is a slightly larger andmore yellow species close to G. pumila. It was foundin Mårås and Skubbhult. Both species are probablywidespread in Sweden, but have been intermixed with
other species in the genus. In the ordination of thejoint datasets from Halland and Denmark and in thecalculation of similarity coefficients between all Nat-Man datasets Galerina pseudomniophila wasincluded under G. mniophila.
Hohenbuehelia auriscalpium is a spathulate agaric,characterized by its pale felty pileus and a numberof microscopical characters. It was found on two largebeech logs at Valaklitt, both in an intermediate stageof decay. The species is not earlier reported fromSweden, but it is possible that some records of H.reniformis refer to the species. Hohenbueheliaauriscalpium is included among the European beechforest indicators by Christensen et al. (2005) andseems to be obligate in the most valuable beech forestreserves in Europe, where it typically grows on largebeech logs. There are however also records fromother tree species and less valuable habitat types andthe indicator value has still to be proven.
Mycena renati (NT), gulfotshätta, is a beautifulMycena characterized by a yellow stipe, a pinkishcap and a distinct alkaloid smell. It was found in sevenstudy sites in Halland, mostly during the summervisit. The species inhabit various types of deaddeciduous wood and is not uncommon in fertiledeciduous forests in SE Denmark. It seems to havean eastern distribution in Scandinavia and theoccurrence in Halland is slightly surprising as it ismissing form similar habitat-types in Jutland,Denmark. Likewise it was not recorded from Belgiumand the Netherlands during the Nat-Man project(Ódor et al 2004a). It is listed as a relict fungi ofprimeval forests in central Poland (£uszyñski 2003),but according to my experience the species isrestricted more by climatical and forest typeconstraints than breaks in dead wood continuity.
Omphalina epichysium, grånavling, is a small greyagaric with a striate pileus and decurrent gills. It wasfound in Kvinnsåsen and Mårås, and occurred quiteabundantly at the latter locality. All records weremade on large, strongly decayed beech logs with richcommunities of saprotrophic agarics. In central andnorthern Sweden it is mostly found on dead wood ofspruce and aspen and is regarded as an indicator ofhabitat quality (Hallingbäck & Aronsson 1998). Itwas included among the indicators of habitat qualityin Danish beech forests by Heilmann-Clausen &Christensen (2000b), but omitted from the Europeanindicator system (Christensen et al. 2005), mostlydue to the rather small and shortlived sporocarps. InDenmark it is only known from one site, SuserupSkov, which is among the very best sites for
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Saproxylic fungi
saproxylic fungi on beech wood in NW Europe(Heilmann-Clausen & Christensen 2004a).
Ossicaulis lignatilis (syn. Clitocybe lignatilis) (NT),vedtrattskivling, is a fleshy, white to pale grey agaricwith a strong mealy smell. During the project it wasfound on a single beech log at Myskebackarna,associated with Hercium coralloides. The sameassociation was noted to be characteristic in virginforests in eastern Slovakia (Heilmann-Clausen &Christensen 2004a). Ossicaulis lignatilis, however,seems to have rather wide ecological amplitude andis known from a large number of deciduous hosts. Itoccurs in all parts of Sweden and is even known fromsubalpine birch forests (Artdatabanken 2000a). Inbeech forests it seem to be more or less restricted tohabitats of high conservation value and it is includedin the indicator systems of Heilmann-Clausen &Christensen (2000b), Ainsworth (2004) andChristensen et al. (2005). Ossicaulis lignatilis is oneof the rather few agarics that cause brown rot and itis most likely a heart rot agent, associated mainlywith old, overmature trees.
Pluteus umbrosus (NT), borstskölding, is a highlycharacteristic species, identified by the flesh-colouredgills with a dark edge and the beautiful velvety browncap surface. The species belongs to a characteristicsociety of agarics growing on deciduous wood in a
rather advanced stage of decay. It is found on deadwood of several deciduous tree species, of whichbeech, birch and aspen appear to be most commonin Sweden (Hallingbäck & Aronsson 1998). It wasfound in seven of the study sites (Fig. 13) and wasslightly more common in Halland, than in any of theNat-Man datasets (cf Ódor et al. 2004a). The speciesis apparently a good indicator of sites with a stablerepresentation of well decayed deciduous wood, andis included in the indicator systems of both Heilmann-Clausen & Christensen (2000b), Nitare (2000) andChristensen et al. (2005).
Psathyrella larga is a rather large Psathyrella whichis here reported as new to Sweden, thanks todetermination work by Leif Örstadius, Kristiansstad.The species is rather widely distributed in Sweden,but has previously been confused with similar lookingspecies like P. obtusata and P. rostellata. It is evencollected in Germany, Denmark, Norway andFinland. It has no special connection to beech forests,but occurs in most forest types on soil and wood. Inthe present study it was found in two localities; NKroksjön and Frodeparken, in both cases on stronglydecayed beech wood. (All data on ecology anddistribution are due to Leif Örstadius pers. comm.).
Volvariella bombycina (NT), silkesslidskivling, is alarge, fleshy agaric, which grows in dense clusters
Hohenbeuhelia auriscalpium is recognized on account of the pale, tough, slightly felty and spathuliformsporocarps. Valaklitt. Photo: Jacob Heilmann-Clausen.
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Halland 2004
on dead deciduous wood. The cream-coloured, silkyfibrillose cap and the volvate stem base makes thespecies unmistakeable. It was found in a cavity on aliving beech in Holkåsen, but has also been recordedin a similar situation from N Kroksjön by Örjan Fritz(pers. comm.). The species has a southern to easterndistribution in Scandinavia and is mostly recordedin regions with high summer temperatures. It growson several deciduous tree species and is mostcommon on cavities in still living, somewhat sun-exposed trees, but is also recorded from fallen logsand dead snags. In Halland it seems to depend on awarm, local microclimate. Volvariella bombycina isred-listed also in Denmark (Stoltze & Pihl 1998) andNorway (Bendiksen et al. 1997), and included in thebeech forest indicator systems of both Heilmann-Clausen & Christensen (2000b), Ainsworth (2004)and Christensen et al. (2005).
Hypoxylon cohaerens (NT), liten bokdyna, is apyrenomycete forming rather small, adpressed,blackish stromata on dead beech wood. It is far themost frequently recorded red-listed species in theproject with records from almost all study sites (Fig.14). In the intensive localities it was found on 34 of200 study trees. The species grows on rather freshbeech wood, especially on thick branches, and is mostlikely a latent invader being present in a passive,endophytic stage in living trees. It is almost strictly
associated with beech. The species is rather commonin Denmark, but was included among the indicatorsof habitat quality in Danish beech forests byHeilmann-Clausen & Christensen (2000b). It waslater omitted from the European indicator system(Christensen et al. 2005) because of a rather weakassociation with special habitat qualities.
Nemania atropurpurea & N. chestersii arepyrenomyces forming thin but widely effused,blackish stromata on dead wood. Macroscopicallyboth species looks quite like the more well-knownN. serpens, and it requires some field experience todistinguish the three species in the field, with justsome success. Microscopically the species are easilydistinguished (e.g. Læssøe et al. 2000). Both speciesare here reported as new to Halland, but appear to bequite widespread in the county. This is especially thecase for N. atropurpurea which was found on 13study sites and 11 % of the intensive study trees. Thisfrequency is considerably higher than recorded inany of the Nat-Man countries. Nemania atropurpurea& N. chestersii were both included among theindicators of habitat quality in Danish beech forestsby Heilmann-Clausen & Christensen (2000b), butwere omitted from the European indicator system(Christensen et al. 2005) because of the lack of goodmacroscopical characters. In Halland the two specieswere found to be associated with snags decayed by
Pluteus umbrosus (borstskölding), on strongly decayed beech log in Holkåsen. Photo: Jacob Heilmann-Clausen.
52
Saproxylic fungi
Fomes fomentarius, either standing or recent fallen.Especially N. atropurpurea was very characteristicin this habitat.
Camarops tubulina (NT), gransotdyna, is a stromaticpyrenomycete, forming large and thick, blackstromata on dead wood of fir, spruce and beech. Itwas recorded from eight study sites in the currentproject, all of high to very high conservation interest(Fig. 14). In Sweden the species has previously beenrecorded mostly from fertile spruce forests rich indead wood (Artdatabanken 2000b). In Central Europethe species occur in pure beech forests and mixedfir/beech forests and is included in the beech forestindicator systems of both Heilmann-Clausen &Christensen (2000b) and Christensen et al. (2005).According to Holec (2003) the species is restrictedto forests minimally influenced by human activitiesin Böhmerwald, the Czech Republic. In Halland itoccurred in all cases on standing or fallen beechsnags, which seem to be a preferred habitat also inDenmark. As shown in Fig. 17 it seems to have apreference for large diameter trees.
Peziza saniosa (VU), blåmjölkig storskål, is a disco-mycete forming blackish lens-shaped sporocarps(apothecia), exceeding blue “milk” upon breakage.It was recorded in Nissaström as new to Halland.The species usually grows on exposed, rich clay soilsin ditches, hazel-groves etc. More occasionally it isfound on strongly decayed wood as in the presentcase. The species is rare in Sweden with scatteredrecords mainly in the southern part of the country(Gärdenfors 2000).
Ascocoryne sp. Two species of Ascocoryne arecommonly reported from dead wood in Scandinavia,i.e. A. sarcoides and A. cylichnium (e.g. Christensen& Heilmann-Clausen 2000; Hansen & Knudsen2000). Both were recorded in this project, but inaddition a third species, characterized by rather small,brownish sporocarps and distinctly capitateparaphyses, were found in several localities. Materialhas been send to Viktor Kucera in Slovakia which ispresently working on the genus in his PhD project.
Camarops tubulina (gransotdyna) on beech wood in Skrockeberg. Photo: Örjan Fritz.
53
Halland 2004
Appendix 3. Ordination diagrammes
Diagram showing the optima of fungal species in theordination space defined by axis 1 and 2 in the DCAordination of the Halland dataset (Fig. 8). Specieswith optima on slightly decayed tree have high scores
xy lhy p
f omf om
marall
galmar
lacame
stehireutspi
my chae
psapil
panser
bjeadu
plucer
phlrad
pezmic
kredeu
hy pf as
my ccin
sterug
f ompinly cper
hy pcoh
my cgal
asccy l
antsem
armlut
steoch
inopet
hy pf raneopur
my cspe
megpla
calcor
ly cpy r
oudmuc
nemser
ganlip
hy pmulphlf ag
nematr
my ceru
lenmuc
copmic
datmol
phlliv
plupod
rusmai
ascsar
trav er
lacble
xerrad
phlruf
ricf ib
cantub
plepul
hellac
lacsub
polv ar
nemche
cerexc
my chie
phltre
pluumb
psecy a
hy druf
trahir
hy plat
galv it
camtub
laclac
exigla
lactab
phomut
polbru
climet
my csan
galatk
hy pruf
inonodpsaart
ty rchi
my cpse
schcom
laxbic
possub
strcy a
colbut
pluphl
phy san
tragib
gy msap
hy prubmy cren
consub
corano
galmni
bolret
phaimp
lepf el
pleost
hercor
pansti
cy scar
pluluc
colcoo
entcet
paxinv
peninc
rusoch
hy prad
ly cf oe
phoaur
clihob
clacor
mutcan
my ccro
plupla
crev er
corcas
neccocc
trehy m
cy lev o
dipblox
diadis
skev ul
asc sp
strhor
cy sami
my cmet plusemi
Axis 1
Axi
s 2
on axis 1, while the opposite is true for species withoptima on strongly rotted trees. Species names areabbreviated according to Appendix 4.
54
Saproxylic fungi
xy lhy pf omf om
eutspi
marall
galmar
my chae
kredeu
stehir
plucer
lacame
my cgal
hy pf as
asccy l
psapil
bjeadu
armlut
ly cpy r
panser
copmic
sterug
f ompinphlrad
pezmic
ly cper
ganlip
my ceru
my ccro
hy pcoh
hy pf ra
calcoroudmuc
neopur
phy v itricf ib
antsem
nemser
lacsub
my ccin
phlf ag
steoch
my cspe
ascsar
trav er datmol
psecy a
polv ar
inopet
megpla
hy pmul
nemche
clihob
phlliv
ramstr
consub
pluphl
nematr
rusmai
ty rchi
galv it
plupod
xerrad
phltre
trahir
inonod
lacble
lenmuc
cerexc
colcoomy csan
camtub
exigla
laclac
pluumbclimet
phlrufphomut
my chie
my cmin
plupla
xy lpol
bulinq
cantub
galatk
hy ppar
hy pruf
galhy p
hellac
hy prub
possubplepul
plusal
polbru
galtri
hy plat
pleost
pluluc
plutho
schcomtragib
tremes
chlaer
chopur
hencan
hy druf
hy pmacmarrot
mergig
stesub
hy prad
laxbic
my cren
Skev ul
asc sp
bolret exinuc
lactab
ly cf oe
micbra
my cpsephaimp
psaart
rusoch
simcen
cerret
clidia
cy sami
my cmet
phoaur
strhor
colbut
cordia
cy scar
gy msap
mutcan
psaros
strcy a
auraur
corano
entcet
galmni
hy gaur
iscres
my cpic
paxinv
tref olclacor
cliphy
hercor
lepf el
my cpur
pansti
phef er
phef eu
stef im
campol
humhemlaccam
my cabr
peninc
plurom
psalut
ricswa
bolpru
conbru
consem
corcas
crev er
cy sjas
ganpf e
hohaur
inocin
lenepi
marwy n
melaim
my cf il
my cgau
my cpol
my ctin
ompepi
pholenpsacer
psaobt
resapp
skeniv
trilas
Axis 1
Axi
s 2
Diagram showing the optima of fungal species in theordination space defined by axis 1 and 2 in the DCAordination of the joint datasets from Halland andDenmark (Fig. 9). Species with optima in Hallandand Central Jutland have low scores on axis 2.Species names are abbreviated according to Appendix4, except for the following species, which were notrecorded in Halland: Auricularia auricula-judae(auraur), Camarops polysperma (campol),Chlorociboria aeruginascens (chlaer), Clitocybediatreta (clidia), Conocybe brunnea (conbru), C.semiglobata (consem), Exidia nucleata (exinuc),
Galerina hypnorum (galhyp), Ganoderma pfeifferi(ganpfe), Hypoxylon macrocarpum (hypmac),Ischnoderma resinosum (iscres), Kavinia himantia(kavhim), Marasmius rotula (marrot), M. wynnei(marwyn), Melanophyllum aimatospermum(melaim), Micromphale brassicolens (micbra),Mycena filopes (mycfil), M. galopus (mycgau), M.minutula (mycmin), M. picta (mycpic), Pluteusromellii (plurom), Pluteus thomsonii (plutho),Skeletocutis nivea (skeniv), Steccherinum fimbriatum(stefim) and Tricholoma lascivum (trilas).
55
App
endi
x 4: Lis
t of s
peci
es re
cord
ed in
H
alla
nd
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Asc
omyc
etes
, pyr
enom
ycet
es
Bom
bard
ia a
mbi
gua
var.
carb
onar
ia R
ehm
ss.
Mun
k
0 1
X
C
amar
ops
tubu
lina
(Alb
. & S
chw
.) S
hear
ca
mtu
b 10
8
4%
4%
8%
12%
4%
X
X
X
D
iatry
pe d
isci
form
is (H
offm
.: Fr
.) Fr
.
3 6
X X
X
X
X
X
D
iatry
pe fl
avov
irens
(Per
s.: F
r.) F
r.
1 3
X
X
X
D
iatry
pe s
tigm
a (H
offm
.: Fr
.) Fr
.
2 3
X
X
X
E
utyp
a sp
inos
a (P
ers.
: Fr.)
Tul
. & C
.Tul
. eu
tspi
69
24
30
%
34%
40
%
33%
30
%
64%
12
%
XX
XX
XX
XX
XX
XX
XX
X X
X
Eut
ypel
la c
f. sc
opar
ia (S
chw
ein.
) Ellis
& E
verh
.
1 2
X
X
E
utyp
ella
qua
tern
ata
(Per
s.: F
r.) R
appa
z
2 5
X
X
X
X
X
H
ypoc
rea
argi
llace
a W
. Phi
llips
& P
low
r.
1 1
2%
Hyp
ocre
a ci
trina
(Per
s.: F
r.) F
r.
1 3
4%
X
X
Hyp
ocre
a pu
lvin
ata
Fuck
el
1
1
2%
H
ypoc
rea
rufa
(Per
s.) F
r. hy
pruf
9
5 8%
8%
7%
10%
4%
Hyp
oxyl
on c
ohae
rens
(Per
s.: F
r.) F
r. hy
pcoh
34
18
18
%
12%
24
%
13%
10
%
24%
16
%
XX
X
XX
X
XX
XX
X
Hyp
oxyl
on fr
agifo
rme
(Per
s.: F
r.) K
ickx
hy
pfra
28
25
18
%
4%
20%
13
%
20%
20
%
12%
X
XX
XX
XX
XX
XX
XX
XX
X X
X H
ypox
ylon
mul
tifor
me
(Fr.:
Fr.)
Fr.
hypm
ul
22
13
8%
8%
20%
10%
24
%
8%
X
XX
X
X X
X
Hyp
oxyl
on ru
bigi
nosu
m (P
ers.
: Fr.)
Fr.
hypr
ub
6 8
6%
2%
4%
10
%
X
X
X
X
Kre
tzsc
hmar
ia d
eust
a (H
offm
.: Fr
.) P
. Mar
tin.
kred
eu
39
22
22%
18
%
4%
27%
10
%
28%
24
%
X
XX
XX
XX
XX
X
XX
X X
X
Lasi
osph
aeria
hirs
uta
(Fr.:
Fr.)
Ces
. & D
e N
ot.
0
2
X
X
Nec
tria
cocc
inea
(Per
s.: F
r.) F
r.
17
15
10%
2%
8%
13
%
10%
16
%
8%
X
X
X
XX
X
X
X
Nec
tria
pezi
za (T
ode.
: Fr.)
Fr.
1
3
X
X
X
Nec
triop
sis
viol
acea
(J.C
. Sch
mid
t: Fr
.) M
aire
0 1
X
N
eman
ia a
tropu
rpur
ea (
Fr.:
Fr.)
Pouz
ar
nem
atr
21
14
6%
14%
8%
13
%
20%
16
%
4%
X
X
X
XX
X
Nem
ania
che
ster
sii (
J.D
. Rog
ers
& W
halle
y) P
ouza
r ne
mch
e 12
12
4%
4%
16%
16
%
XX
X
X
X
X
X
X
Nem
ania
ser
pens
(Per
s.: F
r.) G
ray
nem
ser
22
15
22%
2%
8%
7%
8%
20%
X
XX
X
XX
XX
X
Xy
laria
hyp
oxyl
on (L
.: Fr
.) G
rev.
xy
lhyp
14
322
66
%
70%
76
%
80%
60
%
80%
72
%
X
XX
XX
XX
XX
XX
X
X X
X
Xyla
ria p
olym
orph
a (P
ers.
: Fr.)
Gre
v.
xylp
oly
1 1
2%
Asc
omyc
etes
, dis
com
ycet
es
Ale
uria
aur
antia
ca (P
ers.
: Fr)
Fuck
el
0
1
X
Asc
ocor
yne
cylic
hniu
m (T
ul.)
Kor
f as
ccyl
33
14
20
%
10%
24
%
20%
10
%
24%
8%
X
X
X
X
XX
X
56
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Asc
ocor
yne
sarc
oide
s (J
acq.
: Fr.)
Gro
ves
& W
ilson
as
csar
18
11
12
%
6%
12%
10%
20
%
X
X
X
X
X X
A
scoc
oryn
e sp
. as
ctas
9
6 8%
12%
10%
4%
X
X
Asc
otre
mel
la fa
gine
a (P
eck)
Sea
ver
2
4 2%
X
10%
X
B
ispo
rella
citr
ina
(Bat
sch.
: Fr.)
Kor
f & S
.E. C
arp.
10
17
4%
2%
4%
7%
12
%
8%
XX
X
XX
XX
X
X X
X
Bul
garia
inqu
inan
s (P
ers.
: Fr.)
Fr.
bulin
q 6
13
2%
4%
4%
X 8%
X
X
XX
X
X
X
X
C
hlor
ocib
oria
aer
ugin
asce
ns (N
yl.)
Kan
ouse
0 1
X
C
laus
seno
myc
es a
trovi
rens
(Per
s.: F
r.) K
orf &
Aba
wi
0
1
X
cf. D
iplo
carp
a bl
oxam
ii (B
erk.
ex
W. P
hilli
ps) S
eave
r
6 4
X
X X
X
H
elve
lla b
ulbo
sa (H
edw
.: Fr
.) K
reis
el
2
3
4%
X
X
H
elve
lla la
cuno
sa A
fzel
.: Fr
. he
llac
13
6 2%
14
%
33
%
X
X
X
H
umar
ia h
emis
phae
rica
(Wig
g.: F
r.) F
ucke
l hu
mhe
m
3 4
X 4%
4%
X
N
eobu
lgar
ia p
ura
(Fr.)
Pet
rak
neop
ur
28
20
10%
10
%
24%
27
%
30%
16
%
4%
XX
X
X
XX
XX
X
X X
X
Om
brop
hila
sp
1
1
X
Pez
iza
mic
ropu
s P
ers.
: Fr.
pezm
ic
43
15
40
%
16%
33
%
40%
20
%
20%
X
X
X
X
XX
XX
X
P
eziz
a sa
nios
a S
chra
d.: F
r.
0 1
X
Bas
idio
myc
etes
, het
erob
asid
iom
ycet
es
Cal
ocer
a co
rnea
(Bat
sch:
Fr.)
Fr.
calc
or
24
18
20%
6%
12
%
20%
4%
16%
X
XX
X
X
XX
XX
X
X X
D
acry
myc
es e
natu
s (B
erk.
& M
.A. C
urtis
) Mas
see
1
1 X
Exi
dia
glan
dulo
sa (B
ull:
Fr.)
Fr.
exig
la
10
5 2%
2%
8%
16
%
8%
Myx
ariu
m s
ubhy
alin
um (A
. Pea
rson
) D.A
. Rei
d
1 1
X
P
hleo
gena
fagi
nea
(Fr.:
Fr.)
Lin
k ph
lfag
22
8 8%
20
%
12%
10%
8%
8%
X
S
type
lla s
ubge
latin
osa
(P. K
arst
.) P
. Rob
erts
1 1
X
Tr
emel
la fo
liace
a P
ers.
tre
fol
1 2
10%
X
Tr
emel
la m
esen
teric
a R
etz:
Fr.
trem
es
2 4
2%
4%
X
X
Tu
lasn
ella
vio
lea
(Qué
l.) B
ourd
ot &
Gal
zin
2
2 X
X
Bas
idio
myc
etes
, cor
ticoi
ds
Bot
ryob
asid
ium
aur
eum
Par
mas
to
7
16
X X
X X
X X
X
X
XX
XX
X
X
XX
X
X X
B
otry
obas
idiu
m c
onsp
ersu
m J
. Erik
ss.
6
7 X
X X
X
X
X
X
Bot
ryob
asid
ium
sub
coro
natu
m (H
öhn.
& L
itsch
.) D
onk
0
1
X
Bot
ryoh
ypoc
hnus
sub
isab
elin
us (F
r.) J
. Erik
ss.
0
1
X
57
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Cer
aceo
myc
es s
ulph
urin
us (P
. Kar
st) J
. Erik
ss. &
Ryv
arde
n 0
1
?
X
?
Cho
ndro
ster
eum
pur
pure
um (P
ers.
: Fr.)
Pou
z.
chop
ur
3 4
2%
4%
7%
X
Con
ioph
ora
pute
ana
(Sch
umac
h.: F
r.) K
arst
.
2 2
X
X
C
ylin
drob
asid
ium
evo
lven
s (F
r.: F
r.) J
ülic
h
12
5 10
%
2%
8%
8%
8%
Glo
eohy
poch
nici
um a
nalo
gum
(Bou
rdot
& G
alzi
n) H
jorts
tam
0
2
X
X
Hyp
hode
rma
pube
rum
(Fr.)
Wal
lr.
2
2 X
X
Hyp
hode
rma
setig
urum
(Fr.)
Don
k
2 2
X
X
H
ypho
dont
ia p
arad
oxa
(Sch
rad.
: Fr.)
E.L
ange
r &
Ves
terh
. hy
ppar
1
2 2%
X
Hyp
hodo
ntia
radu
la (P
ers.
: Fr.)
E.L
ange
r & V
este
rh.
hypr
ad
4 3
6%
4%
X
H
ypoc
hnic
ium
erik
soni
i Hal
lenb
. & H
jorts
tam
s.l
2
2
X X
Laxi
text
um b
icol
or (P
ers.
: Fr.)
Len
tz
laxb
ic
7 10
6%
2%
8%
4%
X
XX
X
XX
Leuc
ogyr
opha
na s
p
1 2
X
X
M
ycoa
cia
cf. a
urea
(Fr.)
J. E
rikss
. & R
yvar
den.
2 2
4%
7%
P
enio
phor
a ci
nere
a (P
ers.
: Fr.)
Coo
ke
2
2 2%
4%
Pen
ioph
ora
inca
rnat
a (P
ers.
: Fr.)
Kar
st.
peni
nc
4 4
4%
7%
10%
X
P
hane
roch
aete
fila
men
tosa
(Ber
k. &
M. A
. Cur
tis) B
urds
. 1
5 X
X
X
X
X
Pha
nero
chae
te v
elut
ina
(DC
.: Fr
.) K
arst
.
7 5
X X
X
X
X
P
hleb
ia li
vida
(Per
s.: F
r.) B
res.
ph
lliv
20
11
12%
4%
12
%
13%
16%
12
%
X
X
X
X
X
Phl
ebia
radi
ata
Fr.:
Fr.
phlra
d 44
15
14
%
24%
28
%
40%
10
%
28%
16
%
XX
X
XX
XX
X
Phl
ebia
rufa
(Per
s.: F
r.) M
.P.C
hris
t. ph
lruf
14
6 12
%
4%
13
%
8%
8%
X
P
hleb
ia tr
emel
losa
(Sch
rad.
: Fr.)
Bur
ds. &
Nak
as.
phltr
e 11
13
2%
2%
12
%
20
%
16%
X
XX
X
X
X
X X
Sis
totre
ma
radu
loid
es (K
arst
.) D
onk
1
1
X
Sis
totre
ma
sern
ande
ri (L
itsch
.) D
onk
3
3
X
X
X
Sis
totre
mas
trum
niv
eocr
emeu
m (H
öhn.
& L
itsch
.) J.
Erik
ss.
1 1
X
S
tecc
herin
um o
chra
ceum
(Per
s.: F
r..) G
ray
steo
ch
30
14
24%
6%
8%
20
%
X 24
%
16%
X
X
XX
X
X
X
S
tere
um h
irsut
um (W
illd.:
Fr.)
Gra
y st
ehir
71
25
22%
42
%
40%
33
%
40%
60
%
20%
X
XX
XX
XX
XX
XX
XX
XX
X X
X S
tere
um ru
gosu
m (P
ers.
: Fr.)
Fr.
ster
ug
36
20
22%
24
%
16%
13
%
10%
20
%
4%
XX
XX
X
XX
XX
X X
X X
Ste
reum
sub
tom
ento
sum
Pou
z.
stes
ub
2 3
X
4%
4%
Tr
echi
spor
a fa
rinac
ea (P
ers.
: Fr.)
Lib
erta
1 1
X
Tr
echi
spor
a hy
men
ocys
tis (B
erk.
& B
room
e) K
.H. L
arss
. 14
11
X
X
X X
X X
X
X
X
X
X
Trec
hisp
ora
nive
a (p
ers.
) K.H
. Lar
ss.
3
5 X
X X
X
X
58
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Bas
idio
myc
etes
, pol
ypor
es
Ant
rodi
ella
sem
isup
ina
(Ber
k. &
Cur
t.) R
yvar
den
&
Joha
ns.
ants
em
32
16
30%
12
%
12%
13
%
24
%
X
XX
X
XX
X
XX
X
X
Bje
rkan
dera
adu
sta
(Willd
.: Fr
.) K
arst
. bj
eadu
46
19
24
%
18%
36
%
27%
20
%
20%
20
%
XX
XX
XX
XX
XX
X
X
Cer
ipor
ia e
xcel
sa (L
und.
) Par
m.
cere
xc
12
8 8%
2%
8%
20
%
8%
X
X X
Cer
ipor
ia re
ticul
ata
(Hof
fm.:
Fr.)
Dom
. ce
rret
2 3
2%
4%
X
D
aeda
leop
sis
conf
rago
sa (B
olt.:
Fr.)
Sch
roet
.
0 1
X
D
atro
nia
mol
lis (S
omm
erf.:
Fr.)
Don
k da
tmol
20
13
8%
12
%
24%
7%
10
%
4%
4%
X
X
X
X
X X
Dip
lom
itopo
rus
lindb
ladi
i (B
erk.
) Gilb
. & R
yvar
den
0
1
X
Fom
es fo
men
tariu
s (L
.: Fr
.) Fr
. fo
mfo
m
122
25
68%
54
%
64%
53
%
70%
64
%
56%
X
XX
XX
XX
XX
XX
XX
XX
X X
X Fo
mito
psis
pin
icol
a (S
war
tz: F
r.) K
arst
. fo
mpi
n 35
22
36
%
10%
12
%
27%
X
12%
8%
XX
XX
X
XX
XX
XX
X
X X
X G
anod
erm
a lip
sien
sis
(Bat
sch)
Atk
. ga
nlip
22
14
22
%
12%
16
%
X
X 4%
X
X
X
XX
X
XX
H
apal
opilu
s ru
tilan
s (P
ers.
: Fr.)
Kar
st.
0
1
X
Inon
otus
cut
icul
aris
(Bul
l.: F
r.) K
arst
.
0 1
X
In
onot
us n
odul
osus
(Fr.)
Kar
st.
inon
od
8 16
4%
8%
7%
8%
4%
X
X
XX
X
XX
X
X
X
In
onot
us o
bliq
uus
(Per
s.: F
r.) P
ilat
0
3
X
X
X
Lenz
ites
betu
linus
(L.:
Fr.)
Fr.
1
1
2%
M
erip
ilus
giga
nteu
s (P
ers.
: Fr.)
Kar
st.
0
1
X
Oxy
poru
s co
rtico
la (F
r.) R
yvar
den
1
1
10
%
Oxy
poru
s po
pulin
us (S
chum
.: Fr
.) D
onk
0
5
X
X
X
X
X
Phe
llinus
ferre
us (P
ers.
) Bou
rd. &
Gal
z.
phef
er
2 4
4%
X
X
X
Phe
llinus
ferru
gino
sus
(Sch
rad.
: Fr.)
Pat
. ph
efeu
0
1
X
Phy
sisp
orin
us s
angu
inol
entu
s (A
lb. &
Sch
w.:
Fr.)
Pila
t ph
ysan
6
12
4%
4%
10%
8%
XX
X
XX
X
X
X
Phy
sisp
orin
us v
itreu
s (P
ers.
: Fr.)
Kar
st.
2
3 2%
4%
X
Pol
ypor
us b
rum
alis
(Per
s.) F
r.: F
r. po
lbru
9
8
10%
X
7%
10%
8%
X
X
X
Pol
ypor
us c
iliat
us F
r.: F
r.
0 1
P
olyp
orus
var
ius
(Per
s.) F
r.: F
r. po
lvar
12
13
8%
6%
7%
20%
8%
X
XX
X
X
X
X
X
Pos
tia s
ubca
esia
(Dav
id) J
ülic
h po
ssub
7
10
6%
2%
8%
4%
X
XX
X
X
X
P
ostia
teph
role
uca
(Fr.:
Fr.)
Jül
ich
0
1
X
Pyc
nopo
rus
cinn
abar
inus
(Jac
q.: F
r.) K
arst
.
0 1
X
S
kele
tocu
tis v
ulga
ris (F
r.:) N
iem
elä
& Y
.C. D
ai
Ske
vul
10
10
6%
X
10
%
20%
4%
X
X
X
X
X
Tr
amet
es g
ibbo
sa (P
ers.
: Fr.)
Fr.
tragi
b 6
9 6%
7%
10
%
4%
X
X
X
X
X
59
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Tram
etes
hirs
uta
(Wul
fen:
Fr.)
Pila
t tra
hir
10
17
4%
16
%
10
%
12%
XX
X
X
XX
XX
XX
X
X X
Tram
etes
ver
sico
lor (
L.: F
r.) Q
uel.
trave
r 17
16
4%
10
%
24%
7%
20
%
4%
X
XX
XX
X
X
XX
X
Ty
rom
yces
chi
oneu
s (F
r.: F
r.) K
arst
. ty
rchi
8
12
4%
2%
12%
X
8%
X
XX
XX
X
X
Tyro
myc
es s
p.
0
4
X
X
X X
Bas
idio
myc
etes
, oth
er a
phyl
loph
oral
es a
nd
gast
rom
ycet
es
Can
thar
ellu
s tu
baef
orm
is (B
ull.:
Fr.)
Fr.
cant
ub
13
9
6%
8%
20%
20
%
X
X
XX
X
Cla
vulin
a co
rallo
ides
(L. :
Fr.)
J. S
chro
et.
clac
or
3 5
4%
7%
4%
X
X
Den
tipel
lis fr
agilis
(Per
s.: F
r.) D
onk
denf
ra
4 7
4%
8%
X
X
X
X
X
H
enni
ngso
myc
es c
andi
dus
(Per
s.: F
r.) O
.K.
henc
an
1 1
2%
Her
iciu
m c
oral
loid
es (S
cop.
: Fr.)
Per
s.
herc
or
4 11
4%
X
X 8%
XX
XX
X
X
Lent
aria
epi
chno
a (F
r.) C
orne
r le
nepi
0
1
X
Lyco
perd
on fo
etid
um B
onor
d.
lycf
oe
4 1
8%
Lyco
perd
on p
erla
tum
Per
s.: P
ers.
ly
cper
35
18
8%
24
%
16%
40%
16
%
28%
X
XX
X
XX
XX
X
X
X X
Ly
cope
rdon
pyr
iform
e Sc
haef
f.: P
ers.
ly
cpyr
24
16
32%
12
%
10
%
8%
8%
XX
XX
XX
XX
X
X X
Muc
rone
lla c
alva
(Alb
. & S
chw
ein.
) Fr.
1
1
X
Mul
ticla
vula
muc
ida
(Per
s.: F
r.) R
.H. P
eter
sen
mul
muc
21
9
30
%
16%
7%
10
%
X
X
X
X
Mut
inus
can
inus
(Hud
s.: P
ers.
) Fr.
mut
can
3 5
7%
10
%
4%
X
X
P
hallu
s im
pudi
cus
L.: P
ers.
ph
aim
p 5
6 2%
4%
4%
4%
X
X
Ram
aria
stri
cta
(Per
s.: F
r.) Q
uél.
2
5
X
4%
4%
X
X
Scl
erod
erm
a ci
trinu
m P
ers.
1 1
4%
Stig
mat
olem
ma
urce
olat
a (W
allr.
: Fr.)
Don
k
1 1
2%
Bas
idio
myc
etes
, aga
rics
and
bole
tes
Aga
ricus
dul
cidu
lus
S. S
chul
z. in
Kal
kbr.
0
1
X
Aga
ricus
syl
vico
la (V
itt.)
Pec
k
0 1
X
Am
anita
citr
ina
(Sch
eaff.
) Per
s.
0
1
X
Arm
illar
ia lu
tea
Gill
et
arm
lut
31
19
6%
24%
4%
20
%
X 40
%
8%
XX
X
XX
XX
XX
X
X
X
Bol
bitiu
s re
ticul
atus
(Per
s.: F
r.) R
ick.
bo
lret
5 6
2%
7%
10%
8%
X
X
Bol
etus
pas
cuus
(Per
s.) K
rom
bh.
2
2
8%
X
B
olet
us p
ruin
atus
Fr.
2
3 2%
4%
X
Clit
ocyb
e m
etac
hroa
(Fr.:
Fr.)
Kum
m.
clim
et
9 7
12
%
10%
8%
X
X
X
X
60
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Clit
ocyb
e od
ora
(Bul
l.: F
r.) K
umm
.
1 3
2%
X
X
Clit
ocyb
e ph
yllo
phila
(Per
s.: F
r.) K
umm
. cl
iphy
1
2
2%
X
C
litop
ilus
hobs
onii
(Ber
k.) P
.D.O
rton
clih
ob
3 6
4%
4%
X
X
X
X
C
olly
bia
cook
ei (B
res.
) J.D
.Arn
old
colc
oo
4 6
12
%
4%
X
X
X X
Col
lybi
a fa
giph
ilus
(Vel
en.)
Ant
onín
, Hal
ling
& N
oord
el.
2
2
4%
4%
Con
ocyb
e su
bpub
esce
ns P
.D.O
rton
cons
ub
6 7
4%
4%
10%
4%
X
X
X
Cop
rinus
lago
pus
(Fr.:
Fr.)
Fr.
1
1
10
%
Cop
rinus
mic
aceu
s (B
ull.:
Fr.)
Fr.
copm
ic
20
14
8%
14%
4%
7%
30
%
12%
4%
X
X
XX
X
XX
C
oprin
us s
pela
ioph
ilus
Bas
& U
ljé
0
1
X
Cor
tinar
ius
(tela
mon
ia) s
p. 1
1 1
4%
Cor
tinar
ius
(tela
mon
ia) s
p. 2
1 1
4%
Cor
tinar
ius
anom
alus
(Fr.:
Fr.)
Fr.
cora
no
6 4
2%
4%
4%
12
%
Cor
tinar
ius
casi
miri
(Vel
en.)
Hui
jsm
an
corc
as
3 2
13
%
4%
Cor
tinar
ius
cf. a
lbov
arie
gatu
s (V
elen
.) M
elot
1 1
7%
Cor
tinar
ius
deci
pien
s (P
ers.
: Fr.)
Fr.
2
1
8%
C
ortin
ariu
s di
asem
ospe
rmus
D. L
am.
2
2
4%
4%
Cre
pido
tus
appl
anat
us (P
ers.
) Kum
m.
0
2
X
X
Cre
pido
tus
vers
utus
(Pec
k) S
acc.
cr
ever
3
3
2%
8%
X
Cys
tode
rma
amia
nthi
num
(Sco
p.) K
on. &
Mau
bl.
cysa
mi
8 9
10
%
10%
4%
4%
X
X
X
X
X
C
ysto
derm
a ca
rcha
rias
(Per
s.) K
onr.
& M
aubl
. cy
scar
4
8 2%
2%
8%
X
X
X
X
X
C
ysto
derm
a ja
soni
s (C
ooke
& M
ass.
) Har
m.
2
3
7%
4%
X
Ent
olom
a al
boto
men
tosu
m N
oord
el. &
Hau
skn.
1 1
2%
Ent
olom
a ce
tratu
m (F
r.: F
r.) M
oser
en
tcet
4
5
4%
4%
4%
X
X
E
ntol
oma
conf
eren
dum
(Brit
z.) N
oord
el.
2
2 2%
4%
Ent
olom
a pl
acid
um (F
r.: F
r. ) N
oord
el.
1
2 2%
X
E
ntol
oma
prot
erum
Noo
rdel
. & W
ölfe
l
1 1
2%
Ent
olom
a rh
odop
oliu
m (F
r.: F
r.) K
umm
.
0 1
X
G
aler
ina
atki
nson
iana
A.H
.Sm
ith
gala
tk
9 6
2%
8%
4%
7%
10%
4%
Gal
erin
a cf
. lae
vis
(Per
s.) S
ing.
0 1
X
G
aler
ina
mar
gina
ta (B
atsc
h) K
ühne
r ga
lmar
92
20
28
%
56%
44
%
47%
70
%
56%
44
%
XX
X
XX
XX
XX
X
X X
X
Gal
erin
a pa
llida
(Pilá
t) H
orak
& M
oser
0 1
X
G
aler
ina
pseu
doba
dipe
s Jo
ss.
0
1
X
61
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Gal
erin
a ps
eudo
mni
ophi
la K
ühne
r ga
lmni
7
3 8%
2%
10
%
Gal
erin
a sa
hler
i (Q
uel.)
Küh
n.
0
1
X
Gal
erin
a tri
scop
a (F
r.) K
ühne
r ga
ltri
1 1
4%
Gal
erin
a vi
ttaef
orm
is (F
r.) S
inge
r ga
lvit
10
9 2%
8%
8%
8%
4%
X
X
X
X
G
aler
inia
vex
ans
Smith
& S
ing.
0 2
X
X
G
ymno
pilu
s sa
pine
us (F
r.: F
r.) M
aire
gy
msa
p 6
9
2%
10%
12
%
4%
X
X
X
X
X
Heb
elom
a cr
ustu
linifo
rme
(Bul
l.) Q
uél.
1
1
4%
H
ohen
bueh
elia
aur
isca
lpiu
m (M
aire
) Sin
ger
2
1
8%
H
ohen
bueh
elia
flux
ilis (F
r.: F
r.) P
.D. O
rton
1
1
7%
H
ydnu
m ru
fesc
ens
Fr.
hydr
uf
11
3
8%
20
%
24%
H
ygro
phor
opsi
s au
rant
iaca
(With
.: Fr
.) M
aire
hy
gaur
1
1
10
%
Hyp
holo
ma
fasc
icul
are
(Hud
s.: F
r.) P
.Kum
m.
hypf
as
39
21
8%
18%
32
%
13%
10
%
32%
28
%
XX
XX
XX
XX
XX
XX
X
X
Hyp
holo
ma
late
ritiu
m (S
chae
ff.: F
r.) P
.Kum
m.
hypl
at
10
16
2%
8%
4%
7%
10%
4%
4%
X
XX
XX
X
XX
X
H
ypho
lom
a m
argi
natu
m (P
ers.
: Fr.)
Sch
roet
.
2 4
7%
4%
X
X
H
ypsi
zygu
s te
ssul
atus
(Bul
l.: F
r.) S
ing.
0 1
X
In
ocyb
e ci
ncin
nata
(Fr.:
Fr.)
Qué
l.
1 1
4%
Inoc
ybe
petig
inos
a (F
r.: F
r.) G
illet
inop
et
29
14
2%
18%
32
%
27%
10
%
12%
12
%
XX
X
X
X
X
X
Inoc
ybe
sind
onia
(Fr.)
Kar
st.
1
1
4%
La
ccar
ia a
met
hyst
ina
Coo
ke
laca
me
75
19
24%
44
%
32%
47
%
50%
40
%
44%
X
XX
XX
XX
XX
XX
X
La
ccar
ia la
ccat
a (S
cop.
: Fr.)
Ber
k. &
Br.
lacl
ac
10
6 12
%
4%
4%
4%
X
X
La
ctar
ius
blen
nius
(Fr.:
Fr.)
Fr.
lacb
le
17
12
8%
8%
7%
4%
36%
X
X
X
X
XX
X
La
ctar
ius
cam
phor
atus
(Bul
l.: F
r.) F
r. la
ccam
3
2
4%
8%
Lact
ariu
s su
bdul
cis
(Bul
l.: F
r.) G
ray
lacs
ub
13
6 8%
13
%
24
%
4%
X
X
Lact
ariu
s ta
bidu
s Fr
. la
ctab
9
10
4%
16
%
7%
10%
4%
XX
X
X
X
Lent
inel
lus
coch
leat
us (P
ers.
: Fr.)
Kar
st.
0
1
X
Lent
inel
lus
ursi
nus
(Fr.)
Küh
n.
0
2
X
X
Lent
inel
lus
vulp
inus
(Sow
erby
: Fr.)
Küh
n. &
Mai
re
0
2
X
X
Lepi
ota
felin
a (P
ers.
) Kar
st.
lepf
el
5 5
8%
4%
X
XX
Le
pist
a nu
da (B
ull.:
Fr.)
Coo
ke
0
1
X
Mar
asm
iellu
s ra
mea
lis (B
ull.:
Fr.)
Sin
ger
0
1
X
Mar
asm
ius
allia
ceus
(Jac
q.: F
r.) F
r. m
aral
l 99
21
58
%
54%
40
%
33%
20
%
48%
56
%
XX
X
XX
XX
XX
X
X X
X X
Meg
acol
lybi
a pl
atyp
hylla
(Per
s.: F
r.) K
otl.
& P
ouz.
m
egpl
a 26
19
10
%
14%
12
%
7%
40%
12
%
12%
XX
XX
XX
XX
X
X
X
X
62
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Myc
ena
abra
msi
i (M
urr.)
Mur
r. m
ycab
r 2
2
2%
4%
Myc
ena
cine
rella
(Kar
st.)
Kar
st.
myc
cin
39
9 14
%
24%
24
%
20%
20
%
24%
12
%
X
X
Myc
ena
croc
ata
(Sch
rad.
: Fr.)
Kum
m.
myc
cro
3 4
6%
X
X
X
Myc
ena
epip
tery
gia
(Sco
p.: F
r.) S
.F.G
ray.
1 2
2%
X
Myc
ena
erub
esce
ns H
öhn.
m
ycer
u 21
12
2%
22
%
16%
7%
30
%
4%
X
X
X
X
X
X
M
ycen
a ga
leric
ulat
a (S
cop.
: Fr.)
Qué
l. m
ycga
l 33
19
8%
16
%
28%
27
%
20%
24
%
8%
XX
XX
XX
XX
X
X X
X
Myc
ena
galo
pus
(Per
s.: F
r.) K
umm
.
0 1
X
M
ycen
a ha
emat
opus
(Per
s.: F
r.) K
umm
. m
ycha
e 66
19
22
%
32%
24
%
33%
50
%
48%
44
%
XX
X
XX
XX
XX
X
X X
Myc
ena
hiem
alis
(Osb
.: Fr
.) Q
uél.
myc
hie
12
6
16%
13%
8%
X
X
X
M
ycen
a m
etat
a (F
r.) K
umm
. m
ycm
et
8 7
2%
8%
4%
7%
10%
X
X
M
ycen
a m
irata
(Pec
k.) S
acc.
1 1
7%
Myc
ena
poly
gram
ma
(Bul
l.: F
r.) G
ray
1
3 2%
X
X
M
ycen
a ps
eudo
corti
cola
Küh
n.
myc
pse
8 5
2%
8%
13
%
8%
4%
M
ycen
a pu
ra (P
ers.
: Fr.)
Kum
m.
myc
pur
3 2
4%
2%
M
ycen
a re
nati
Qué
l. m
ycre
n 6
7 2%
2%
8%
7%
4%
X
X
M
ycen
a sa
ngui
nole
nta
(Alb
. & S
chw
.: Fr
.) K
umm
. m
ycsa
n 9
6 2%
4%
4%
30%
8%
X
Myc
ena
spei
rea
(Fr.:
Fr.)
Gille
t m
ycsp
e 28
10
4%
8%
24
%
27%
20
%
16%
24
%
X
X X
Myc
ena
tintin
abul
um (F
r.) Q
uél.
1
1
2%
M
ycen
a ze
phiru
s (F
r.: F
r.) K
umm
.
1 4
7%
X
X
X
Om
phal
iast
er a
ster
ospo
ra (L
ange
) Lam
oure
1 1
4%
Om
phal
ina
epic
hysi
um (P
ers.
: Fr.)
Qué
l.
1 2
7%
X
Oss
icau
lis li
gnat
ilis (P
ers.
: Fr.)
Red
head
& G
inns
0 1
X
O
udem
ansi
ella
muc
ida
(Sch
rad.
: Fr.)
Höh
n.
oudm
uc
23
18
4%
16%
12
%
13%
24%
8%
X
XX
X
XX
XX
X
X
X X
P
anel
lus
sero
tinus
(Per
s.: F
r.) K
ühn.
pa
nser
47
14
20
%
24%
40
%
20%
30
%
20%
16
%
X
X
X
XX
X
X
Pan
ellu
s st
iptic
us (B
ull.:
Fr.)
Kar
st.
pans
ti 4
8 2%
2%
7%
4%
XX
X
X
Pax
illus
invo
lutu
s (F
r.) F
r. pa
xinv
4
5
4%
4%
4%
X
X
P
holio
ta a
uriv
ellu
s (F
r.) K
umm
. ph
oaur
3
11
4%
4%
X
X
X
X
X
XX
X
X
P
holio
ta le
nta
(Per
s.: F
r.) S
inge
r
1 6
2%
X
X
X
X
X
Pho
liota
lubr
ica
(Per
s.: F
r.) S
ing.
1 1
2%
Pho
liota
mut
abilis
(Sco
p.: F
r.) K
umm
. ph
omut
9
10
4%
10%
7%
X
4%
X
XX
X
X
Pho
liota
squ
arro
sa (W
eige
l: Fr
.) Ku
mm
.
0 1
X
P
leur
ocyb
ella
por
rigen
s (P
ers.
: Fr.)
Sin
ger
0
1
X
63
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Ple
urot
us d
ryin
us (P
ers.
: Fr.)
Kum
m.
0
8
X X
X
X
X
X
X
X
Ple
urot
us o
stre
atus
(Jac
q.: F
r.) K
umm
. pl
eost
5
5
4%
X
10%
4%
4%
P
leur
otus
pul
mon
ariu
s (F
r.) Q
uél.
plep
ul
13
8
8%
12%
X
10%
16
%
4%
X
X
Plu
teus
cer
vinu
s (B
atsc
h) S
inge
r pl
ucer
45
19
18
%
26%
20
%
20%
40
%
24%
20
%
XX
X
XX
XX
X
XX
X
X
Plu
teus
luct
uosu
s B
oud.
pl
uluc
4
8
4%
8%
X
X
X
X
X
X
Plu
teus
nan
us (P
ers.
: Fr.)
Kum
m.
0
1
X
Plu
teus
pel
litus
(Per
s.: F
r.) K
umm
.
0 1
X
P
lute
us p
etas
atus
(Fr.)
Gille
t
0 2
X
X
P
lute
us p
hleb
opho
rus
(Ditt
m.:
Fr.)
Kum
m.
plup
hl
7 7
4%
6%
8%
X
X
X
X
Plu
teus
pla
utus
(Wei
nm.)
Gill
et
plup
la
3 4
2%
2%
4%
X
Plu
teus
pod
ospi
leus
Sac
c. &
Cub
. pl
upod
19
7
22
%
8%
16%
8%
X
X
X
P
lute
us s
alic
inus
(Per
s.: F
r.) K
umm
. pl
usal
1
2 2%
X
P
lute
us s
emib
ulbo
sus
(Las
ch) G
ill.
plus
em
4 10
2%
X
4%
7%
10%
X
X
X
XX
P
lute
us u
mbr
osus
(Fr.)
Kum
m.
pluu
mb
11
7
6%
4%
27%
20
%
4%
X
X
P
sath
yrel
la a
rtem
isia
e (P
ass.
) Kon
r. &
Mau
bl.
psaa
rt 8
14
4%
4%
7%
30
%
4%
X
XX
X
X
X
XX
X
P
sath
yrel
la c
ernu
a (V
ahl.:
Fr.)
Hirs
ch
2
3
2%
4%
X
Psa
thyr
ella
cot
onea
(Qué
l.) K
onr.
& M
aubl
.
1 4
4%
X
X
Psa
thyr
ella
fusc
a (S
chum
.) A.
Pea
rs.
1
1
2%
P
sath
yrel
la la
rga
(Kau
ffman
) A.H
. Sm
ith
2
3
2%
4%
X
Psa
thyr
ella
obt
usat
a (P
ers.
: Fr.)
A.H
.Sm
ith
psao
bt
2 3
2%
X
X
Psa
thyr
ella
pilu
lifor
mis
(Bul
l.: F
r.) P
.D.O
rton
psap
il 48
19
20
%
20%
36
%
13%
50
%
40%
8%
X
XX
X
XX
XX
XX
X X
P
sath
yrel
la ro
stel
lata
Örs
tadi
us
psar
os
0 1
X
P
sath
yrel
la s
arco
ceph
ala
(Fr.)
Sin
g.
0
1
X
Pse
udoc
litoc
ybe
cyat
hifo
rmis
(Bul
l.: F
r.) S
inge
r ps
ecya
11
7
12
%
4%
X
16%
XX
X
Res
upin
atus
app
licat
us (B
atsc
h: F
r.) G
ray
1
2
2%
X
R
hodo
colly
bia
buty
race
a (B
ull.:
Fr.)
Len
nox
colb
ut
7 9
6%
6%
10%
X
X
X
XX
X
R
icke
nella
fibu
la (B
ull.:
Fr.)
Rai
th.
ricfib
13
11
6%
6%
4%
13
%
12
%
4%
X
X
X
XX
Ric
kene
lla s
war
tzii
(Fr.:
Fr.)
Kuy
per
ricsw
a 1
1
4%
R
ussu
la fe
llea
(Fr.)
Fr.
1
1
4%
R
ussu
la m
aire
i Sin
ger
rusm
ai
18
14
2%
6%
16%
10%
12
%
24%
X
X
X
XX
X
XX
R
ussu
la o
chro
leuc
a P
ers.
ru
soch
4
4 6%
4%
X
X
Sch
izop
hyllu
m c
omm
une
Fr.:
Fr.
schc
om
7 10
2%
2%
4%
20%
8%
X
X
X
XX
64
Code used in ordination diagrams
number of trees
Number of sites
Dömestorp
Holkåsen
N Kroksjön
Kvinnsåsen
Trälhultet
Valaklitt
NV Äskemossen
NÖ Kroksjön S. Ry Mårås
Getabäcken Blåalt
Almeberget Skubbhult
Frodeparken Myskebackarna
Skrockeberg Svarta Klippan
Råmebo Vallåsen
Klinta hallar Nissaström Rågetaåsen Ödegärdet Djupeåsen
Sim
ocyb
e ce
ntun
culu
s (F
r.: F
r.) K
arst
. si
mce
n 2
4
2%
4%
X
X
Sim
ocyb
e ru
bi (B
erk.
) Sin
ger
2
1
4%
S
phae
robo
lus
stel
latu
s To
de: P
ers.
1 1
4%
Stro
phar
ia a
erug
inos
a (C
urt.:
Fr.)
Qué
l.
0 1
X
S
troph
aria
cya
nea
(Bul
l.) T
uom
ikos
ki
strc
ya
7 10
4%
4%
4%
7%
4%
X
X
X
X
X
S
troph
aria
hor
nem
anni
i (Fr
.: Fr
.) Lu
ndel
l st
rhor
8
12
X 2%
12
%
7%
20%
4%
XX
X
X
X
X
Tylo
pilu
s fe
lleus
(Bul
l.: F
r) K
arst
.
1 1
7%
Vol
varie
lla b
omby
cina
(Sch
aeff.
: Fr.)
Sin
ger
0
1
X
Xeru
la ra
dica
ta (R
elha
n: F
r.) D
örfe
lt xe
rrad
15
16
6%
12%
4%
13
%
30%
X
X X
X
XX
X
X
X
X
X
Steril
e m
ycel
ia
Ant
hini
a fla
mm
ea (J
ungh
.) Fr
.
1 1
X
App
endi
x 4.
Lis
t of a
ll fu
ngal
spec
ies r
ecor
ded
in th
e pr
ojec
t, so
rted
in m
ajor
mor
phol
ogic
al g
roup
s. Th
e lo
catio
ns o
f stu
dy si
tes a
re sh
own
in F
ig. 1
. For
the
inte
nsiv
e st
udy
site
s (C
olum
ns 5
– 1
1) th
e pe
rcen
tage
s of t
rees
with
spor
ocar
ps o
f eac
h sp
ecie
s are
giv
en. A
n “X
” in
dica
te
that
the
spec
ies w
as o
nly
reco
rded
supp
lem
enta
ry o
n tre
es n
ot in
clud
ed in
the
inte
nsiv
e st
udy.
