Magne Sætersdal

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Sammendrag

Data på forekomster av karplanter, lav, moser og vedboende sopp innsamlet i 1997-98 ble analysert med hensyn på artsmangfold og artssammensetning i eldre naturskog og yngre kulturskog i tre ulike studieområder. Det ble funnet klare forskjeller mellom naturskog og kulturskog for lav og sopp (flere arter og annerledes artssammensetning i naturskog), men mindre slike forskjeller for karplanter og moser.

Sammendrag

I rapporten sammenlignes resultatene av to ulike MiS-registreringer gjennomført i henholdsvis 2009 og 2020 i Follsjåområdet i Notodden kommune. Resultatene viste betydelige forskjeller mellom takstene både når det gjaldt totalt areal av registrerte livsmiljøer, fordeling av areal på ulike typer livsmiljøer, og evnen til å fange opp viktige livsmiljøer for rødlistearter. Vi diskuterer årsaker til disse forskjellene, og foreslår også noen muligheter for forbedringer av registreringene.

Sammendrag

I rapporten undersøkes og diskuteres hvilke skogtyper innen det Europeiske Natura 2000 systemet som Norge synes å ha et særlig ansvar for. Data fra Landsskogtakseringen sammenlignes med data fra EU på areal av de ulike skogtypene. Det konkluderes med at Norge har et svært høyt ansvar for oseaniske skogtyper, et høyt ansvar for Nordiske skogtyper, og et middels til lavt ansvar for sydlige skogtyper.

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Rapporten presenterer resultater fra en undersøkelse av arealer registrert som MiS-livsmiljøet Rik bakkevegetasjon i tre områder på Østlandet. De overordnete målene var å undersøke om arealene tilfredstilte kravene til vegetasjonstyper som definerer livsmiljøet, og om de utvalgte arealene satt av som nøkkelbiotoper skilte seg ut fra arealer utenfor nøkkelbiotopene på samme bonitet og skogtype med hensyn på forekomster av arter innen karplanter og jordboende sopp. Prøveflater (50x50m) i tilfeldig utvalgte nøkkelbiotoper inneholdt Rik bakkevegetasjon i 92% av tilfellene, og hadde en høyere andel av kalklågurtskog og høystaudeskog enn referanseflatene. Det ble funnet et høyere antall karplanter i MiSflatene enn i referanseflatene, men ikke jordboende sopp. For sopp på rødlisten ble det funnet et stort antall rødlistearter, et noe høyere antall arter i MiS-flatene enn i referanseflatene, og at fem av seks flater med >10 rødlistete sopp var MiS-flater. Forskjeller i sammensetningen av arter på flatene var i hovedsak knyttet til geografisk beliggenhet og til vegetasjontyper. Flater i middelaldret kulturskog hadde færre karplanter, men minst like mange arter sopp (inkludert rødlistearter) som naturlig forynget skog. Resultatene viser at nøkkelbiotoper basert på MiS-registreringer fanger opp de vegetasjonstypene og arter som definerer livsmiljøet, men at bruk av skillearter innen karplanter for å lokalisere kalklågurtskog ikke alltid fungerer i tett skog med lite karplanter.

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Red-listed species are often used as target species in selection of sites for conservation. However, limitations to their use have been pointed out, and here we address the problem of expected high spatio-temporal dynamics of red-listed species. We used species data (vascular plants, bryophytes, macrolichens and polypore fungi) from two inventories 17 years apart to estimate temporal turnover of red-listed and non-red-listed species in two forest areas (147 and 195 ha) and of plots (0.25 ha) within each area. Furthermore, we investigated how turnover of species afected the rank order of plots regarding richness of red-listed species, using two diferent national Red List issues (1998 and 2015). In both study areas, temporal turnover was substantial, despite minor changes in the overall number of species. At plot level, temporal turnover in red-listed species was higher than in non-red-listed species, but similar to non-red-listed species of the same frequency of occurrence. Adding the efect of changing identities of species red-listed according to the two Red List issues, further increased the estimated spatio-temporal dynamics. Recorded spatio-temporal turnover also resulted in substantial changes in the rank order of plots regarding richness of red-listed species. Using rare red-listed species for site selection may therefore be accompanied by a higher loss of conservation efectiveness over time than for more common species, and particularly at fner scales. Red-listed species · Site selection · Spatio-temporal dynamics · Temporal turnover

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The pine-dominated forests of Western Norway have been found to harbour viable populations of woodpeckers, including the highly specialized White-backed Woodpecker Dendrocopos leucotos. The aim of this study was to investigate to what extent there were any changes in frequencies of woodpeckers, in particular the White-backed Woodpecker and the Grey-headed Woodpecker Picus canus, by resurveying 60 plots (each 1 km2 ) originally surveyed during 1994/1995. The resurvey was performed in 2013/2014. The White-backed Woodpecker was found to be the most common woodpecker species in both time periods. The Grey-headed Woodpecker was found to have a statistically significant decline from 27% of the 60 plots in 1994/95 to only 12% in 2013/14. The other four species all increased in frequency; although none of those increased frequencies were found to be statistically significant. We discuss possible explanations to why pine forests in Western Norway constitute a valuable habitat for the White-backed Woodpecker at the same time as it has drastically declined in other parts of Norway and Western Europe. In general, the reduced frequency of Grey-headed Woodpecker is not fully understood, although we suggest that cold winters during the years prior to the surveys in 2013/14 may be an important factor.

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Aim: Revisits of non-permanent, relocatable plots first surveyed several decades ago offer a direct way to observe vegetation change and form a unique and increasingly used source of information for global change research. Despite the important insights that can be obtained from resurveying these quasi-permanent vegetation plots, their use is prone to both observer and relocation errors. Studying the combined effects of both error types is important since they will play out together in practice and it is yet unknown to what extent observed vegetation changes are influenced by these errors. Methods: We designed a study that mimicked all steps in a resurvey study and that allowed determination of the magnitude of observer errors only vs the joint observer and relocation errors. Communities of vascular plants growing in the understorey of temperate forests were selected as study system. Ten regions in Europe were covered to explore generality across contexts and 50 observers were involved, which deliberately differed in their experience in making vegetation records. Results: The mean geographic distance between plots in the observer+relocation error data set was 24 m. The mean relative difference in species richness in the observer error and the observer+relocation data set was 15% and 21%, respectively. The mean “pseudo-turnover” between the five records at a quasi-permanent plot location was on average 0.21 and 0.35 for the observer error and observer+relocation error data sets, respectively. More detailed analyses of the compositional variation showed that the nestedness and turnover components were of equal importance in the observer data set, whereas turnover was much more important than nestedness in the observer+relocation data set. Interestingly, the differences between the observer and the observer+relocation data sets largely disappeared when looking at temporal change: both the changes in species richness and species composition over time were very similar in these data sets. Conclusions: Our results demonstrate that observer and relocation errors are nonnegligible when resurveying quasi-permanent plots. A careful interpretation of the results of resurvey studies is warranted, especially when changes are assessed based on a low number of plots. We conclude by listing measures that should be taken to maximally increase the precision and the strength of the inferences drawn from vegetation resurveys.

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The long-term success of sites selected for species conservation depends on the persistence of target species. Red List species or threatened species lists are frequently defined as target species, but when Red Lists are updated, their species composition may change. Here we investigate the effects of Red List updates on the long-term robustness of fine-scale site selection. We used records of red-listed species (vascular plants, bryophytes, macrolichens, and polypore fungi) recorded in 1997–1998 in 1058 sample plots (50 × 50 m) from six forest landscapes in Norway, and four consecutive issues of the Norwegian Red List for species (1998, 2006, 2010, 2015). Sites were selected based on the first issue (1998) using both a scoring (“hotspot”) approach and a complementarity approach, and the ability of selected sites to include red-listed species of later issues was measured. In four boreal forests the mean proportion of red-listed species included in selected sites were reduced by18% during the study period, whereas no such effect was found in two hemiboreal forests, where increased clustering of red-listed species in sites compensated for target species changes. Changing target species adds to earlier documented challenges caused by population dynamics, and we suggest that alternatives to using occurrences of target species in site selection should be considered, and particularly at finer spatial scales.

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Sammendrag

I rapporten undersøkes mulighetene for å kunne predikere forekomster av MiS-miljøer basert på enkle landskapsvariabler fra kart. Disse variablene er helning, bonitet, høyde over havet og avstand fra vei. Vi bruker oversikt over registrerte MiS-miljøer (bruttolister) fra skogbruksplaner fra 10 kommuner på sør-og østlandet til å utarbeide prediksjonsmodeller. Disse prediksjonsmodellene blir så testet ut i 10 andre kommuner fra samme region. Resultatene viser at det er stor variasjon mellom kommuner i hvor stor grad modellene klarer å fange opp MiS-miljøer. Videre arbeid må undersøke mulighetene for å utarbeide ulike modeller i ulike regioner med ulike typer landskap.

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The forest landscape across the Nordic and Baltic regions hosts numerous lakes and watercourses, which must be included in forest management. In this study, national policy designs regarding protection zones for surface waters on forest land were reviewed and compared for the Nordic countries, Estonia and Latvia. The focus was how each country regulates protection zones, whether they are voluntary or mandatory, and the rationale behind adopting a low or high degree of prescriptiveness. Iceland and Denmark had a low degree of policy prescriptiveness, whereas Norway, Estonia and Latvia had a high degree of prescriptiveness. Sweden and Finland relied to a large extent on voluntary commitments. The prescribed zone widths within the region ranged from 1 m to 5 km. The results indicated that land-use distribution, forest ownership structure and historical and political legacies have influenced the varying degrees of prescriptiveness in the region.

Sammendrag

I rapporten undersøkes og diskuteres mulighetene for å forenkle MiS-registreringer i kyststrøk med mye krevende terreng i forbindelse med feltregistreringer. Vi benyttet registreringer av MiS i Landsskogtakseringen og hogststatestikk som grunnlag for vurderingene. Forutsetningene for forenklinger vurderes å være særlig gode på Vestlandet, men også i Trøndelag finnes muligheter for å redusere arbeidet i felt samtidig som de viktigste livsmiljøene blir registrert.

Sammendrag

I rapporten gjennomgås noen sentrale temaer med relevans for vurdering av miljøregistreringene i skogbruksplanleggingen så langt. Videre skisseres en fremgangsmåte for kvalitetsbedømming av gjennomførte registreringer basert på kvalitetskrav, arealkrav og dokumentasjonskrav, og til slutt oppsummeres anbefalinger for revisjon og videreføring av MiS i en 7-punkts liste.

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Variable retention harvesting is acknowledged as a cost-effective conservation measure, but previous studies have focused on the environmental value and planning cost. In this study, a model is presented for optimizing harvesting cost using a high resolution map generated from airborne laser scanning data. The harvesting cost optimization model is used to calculate the objective value of different scenarios. By comparing the objective values, better estimates of the opportunity cost of woodland key habitats are found. The model can be used by a forest manager when evaluating what silvicultural treatments to implement or as an input for improving the nature reserve selection problem for woodland key habitats or retention patches. The model was tested on four real-world cases, and the results indicate that terrain transportation costs vary more than reported in the literature and that it may be worthwhile to divide the opportunity cost into its direct and indirect components.

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Sammendrag

Retention of selected trees in clear-felling areas has become an important conservation measure in managed forests. Trees with large size or high age are usually preferred as retention trees. In this paper we investigated whether a single large or several small trees should be left in clear-felling areas to serve as life boats and future habitat for epiphytic species. The focal species were 25 Lobarion epiphytic lichens hosted by aspen (Populus tremula). We analyzed the relationships between: (1) proportion of trees colonized and tree size, (2) number of lichen thalli (lichen bodies) and aspen area, and (3) number of lichen species and aspen area, for 38 forest sites. Mixed effect models and rarefaction analyzes showed that large and small host trees had the same proportion of trees colonized, the same number of thalli, and the same species richness for the same area of aspen bark. This indicates that larger aspens do not have qualities, beyond size, that make them more suitable for Lobarion lichens than smaller sized aspen trees. None of the species, not even the red-listed, showed any tendencies of being dependent on larger aspens, and our results therefore did not support a strategy of retaining only large and old trees for conservation of epiphytic Lobarion lichens. Additionally, young aspens have a longer expected persistence than old aspens. However, old retention trees might be important for other species groups. We therefore recommend a conservational strategy of retaining a mixed selection of small/young and large/old aspens.

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Several studies have recently reported that common species are more important for species richness patterns than rare species. However, most such studies have been based on broad-scale atlas data. We studied the contribution of different species occupancy, i.e. number of plots occupied, to species richness patterns emerging from species data in 50 by 50 m plots within six 140–200 ha forests in Norway. The study included vascular plants, lichens, bryophytes, and polypore fungi. We addressed the following questions: 1) are common species more correlated with species richness than rare species? 2) How do occupancy classes combine at various levels of species richness? 3) Which occupancy class is best in identifying the overall most species-rich sites (hotspots) by sampling? The results showed that rare species were better correlated with species richness than common species when the information content was accounted for, that high species richness was associated with a higher proportion of less frequent species, and that the best occupancy class for local hotspot identification was species present in 10–30% of the plots within a forest. We argue that the observed correlations between overall richness and sub-assembly richness are primarily structured by the combination of the distributions of species richness and species occupancy. Although these distributions result from general ecological processes, they may also be strongly affected by idiosyncratic elements of the individual datasets caused by the specific environmental composition of a study area. Hence, different datasets collected in different areas may lead to different results regarding the relative importance of common versus rare species, and such effects should be expected on both broad and fine spatial scales. Despite these effects, we suggest that infrequent species will tend to be more strongly correlated to species richness at local scales than at broader scales as a result of more right-skewed species-occupancy distributions.

Sammendrag

Biodiversity is assumed to have high value for many people, but the necessary preservation also incurs a cost for the forest owner. Typically, studies of this cost are at the stand level, and hence, not very accurate as the cost may vary even within the stand. In this work, we use a 1m×1m grid, generated from LIDAR data, to estimate the cost for harvesting and forwarding. The method could be utilized to calculate compensation, and to select between key woodland habitats to minimise the cost. In three of four test cases, the main cost was reduced harvested volume, but in one case the key woodland habitat also made the harvesting operations more expensive.

Sammendrag

1. Surrogate species measures of biodiversity (SSB) are used worldwide in conservation prioritisations. We address the important question whether the ideas behind SSB are consistent with current knowledge on distribution patterns of species, as reflected in theories of community assembly. 2. We investigated whether assumptions necessary for successful functioning of SSB (nested species assemblages, cross-taxon congruence, spatio-temporal consistency) were supported by predictions from either niche or neutral community models. 3. We found a general mismatch between ideas behind SSB and ecological community theory, except that SSB based on complementarity may be consistent with niche-based theory when gradients in species composition are strong. 4.  Synthesis and applications. The lack of a necessary scientific foundation may explain the disappointing results of empirical tests of SSB. We argue that site selection should be based on costs and opportunities within complementary environmental/land units, rather than expensive inventories of unfounded surrogate species.

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Sammendrag

Riparian forests (RF) growing along streams, rivers and lakes comprise more than 2% of the forest area in the Nordic countries (considering a 10 m wide zone from the water body). They have special ecological functions in the landscape. They receive water and nutrients from the upslope areas, are important habitats for biodiversity, have large soil carbon stores, but may emit more greenhouse gases (GHG) than the uplands. In this article, we present a review of the environmental services related to water protection, terrestrial biodiversity, carbon storage and greenhouse gas dynamics provided by RF in the Nordic countries. We discuss the benefits and trade-offs when leaving the RF as a buffer against the impacts from upland forest management, in particular the impacts of clear cutting. Forest buffers are effective in protecting water quality and aquatic life, and have positive effects on terrestrial biodiversity, particularly when broader than 40 m, whereas the effect on the greenhouse gas exchange is unclear.

Sammendrag

Effekter av skogbruk på biologisk mangfold er i dag et viktig tema innen skogforskningen. Vi vet en hel del om artenes biologi, og hvilken type skog og habitater de finnes i. Dette har blant annet gitt mulighet for å registrere og ta vare på arealtyper som er spesielt viktige for biologisk mangfold. Det vi vet mindre om er hvordan bestandsskogbruket påvirker artenes populasjoner over tid.

Sammendrag

En satsing på kystskogbruket inklusive økt avvirkning de kommende tiår må foregå innenfor faglig funderte rammer som ivaretar sektoransvaret. Hensynet til biologisk mangfold setter ulike rammer og begrensninger for utnyttelsesmulighetene. De praktiske tiltakene skjer på tre fronter: Gjennom vern av skog (etter naturvernloven), gjennom avsetting av viktige områder for biologisk mangfold og de mer generelle hensyn i skogsektoren inkludert gjennom sertifisering og miljøstandarder samt lover og forskrifter. Noen miljøspørsmål som bør ha forskningsmessig fokus de kommende år er gitt korte kommentarer.

Sammendrag

Siden 2001 har det så langt blitt utført miljøregistreringer i forbindelse med skogbruksplanlegging på ca. 65 prosent av alt produktivt skogareal i Norge. Registreringene gjøres etter en metodikk som ble utviklet i prosjektet ”Miljøregistrering i Skog”. Miljøregistreringene gir kartfestet informasjon om forekomst av ulike viktige livsmiljøer i produktiv skog. Bak den praktiske metodikken ligger utarbeidelsen av en generell modell for registrering av biologisk viktige områder. Denne modellen er nå publisert internasjonalt og de viktigste prinsippene er beskrevet her.

Sammendrag

Forest stands are the basic planning units of managed forest landscapes, and the structural composition of these units is important for conservation of biodiversity. We present a methodological approach for identification and mapping of important structural and environmental features of forest stands. Based on an analysis of habitats of red-listed species and a synthesis of results from research on spatial distribution of forest species, we developed a habitat inventory approach (Complementary Hotspot Inventory, CHI) that is currently used in forestry planning in Norway. The CHI maps fine-scale hotspots for 12 habitat types that are further classified according to positions along main environmental gradients (productivity and humidity). Consisting of different substrates in different environments, these habitats to a large degree support different species assemblages. By incorporating both the hotspot and the complementary approach, the CHI produces data tuned for later conservation measures. The high spatial resolution of data facilitates the use of conservation measures at different spatial scales, from single-tree retention to forest reserves. Avalidation test of habitats identified by CHI showed that the density of red-listed species was four times that of randomly selected old forests.

Sammendrag

Results from a literature review on pinewood ecology, silviculture, genetics, aspects of history and forest resources of Scots pine (Pinus sylvestris L.) in western Norway are presented. The pinewoods cover 40 per cent of the forested land, 0.31 million ha. During the last 75 years, the area has increased by 17 per cent and the growing stock has risen from 10 to 34 million m3. The impact of man in previous times was very marked, and has had a significant influence on the present forest conditions. The pronounced climatic gradients mixed with the topographic variation - from the coastal plains via the fjord systems to the high mountains - is reflected in rather steep gradients in the pine forest vegetation. Various floristic elements can be distinguished, from oceanic via the suboceanic in the outer islands to the thermophytic, boreonemoral and boreal elements in the inner fjord districts and valleys. The introduction of spruce (Picea spp.) plantations on 10-15 per cent of former native pine forests has not negatively affected the bird fauna at the landscape scale. Although not particular species rich, the pine forests harbour species usually not found in other forest types. So far, most work in the field of silviculture and forest ecology in the pinewoods of West Norway has been in the form of case studies. Implications of the results for forestry in the region are briefly discussed.

Sammendrag

The potential as indicators of species richness were investigated for 178 species belonging to six ecologically defined species groups (epiphytic bryophytes on nutrient-rich bark, epiphytic macrolichens on nutrient rich bark, pendant lichens on conifer trees, bryophytes on siliceous rocks, bryophytes on dead conifer wood, and polypore fungi on dead conifer wood), using species data from 0.25 ha plots from three different coniferous forest areas (ca. 200 ha each). A species was defined as a potential indicator species for a species group within a study area if its distribution was statistically significantly nested within the species-plot matrix ranked according to species richness, and if the plot frequency of the species was less than 25%. Only two species were identified as potential indicators within all three areas and on average ≈80% of the potential indicator species were lost from one area to another. The results indicate that inconsistency between areas in the species’ frequency distributions and their position in nested hierarchies may strongly reduce the general predictive power of indicator species of species richness, even if significantly nested patterns are found at the community level. We suggest that indicators related to amount and quality of habitats may be an alternative to lists of indicator species of species richness.

Sammendrag

Forest ecosystems provide many deliverables or benefits to society. The most obvious one is wood for the forest industry. Other benefits include berries, hunting, and recreation. More recently recognised benefits are environmental services such as carbon sequestration, water protection and biodiversity, which are without an immediate market value.On the other hand, there are pressures (e.g. climate change, air pollution, exploitation, and costs) on the ecosystem that may hamper the wood production or other benefits......

Sammendrag

Using information from a regional survey of vascular plants of 130 sites in western Norway, a selection of sites based on a heuristic iterative complementarity-based nature reserve selection procedure was performed. The results indicate that conservation of traditionally managed hay meadows is of major importance as they contributed 60.1% of all native species recorded; afforested grasslands (deciduous woodlands < 70 years old) contributed 26.8%, whereas artificially fertilized hay meadows and intensively cultivated grasslands taken together contributed 13.1% of the species. The species composition of the meadows was significantly nested. Thus, if you conserve the most species-rich meadows, you also conserve most of the species in the less species-rich meadows. Nestedness in meadows was significantly correlated with within-meadow habitat diversity and soil pH. The most species-rich meadows were traditional meadows, characterized by high habitat diversity and high soil pH. These meadows will support nearly all species including habitat specialists and regionally rare species, whilst artificially fertilized hay meadows only support the generalist subset, i.e. common species. Area was not significantly correlated with nestedness suggesting that it is more important to cover many habitats than to preserve large traditional meadows just because they are large.

Sammendrag

Vascular plants were investigated as a potential surrogate group in complementary small scale site selection, such as woodland key habitats in Scandinavia. We compared the response of vascular plants to environmental gradients to that of seven other plant, fungal and animal groups within a forest reserve in western Norway using data from 59 plots of 0.25 ha. We also examined whether the spatial changes in species (beta-2 index) of vascular plants matched that of the other groups. All seven groups responded to the same gradients in nutrient richness and humidity as the vascular plants. Furthermore, changes in species composition of vascular plants were reflected in comparable degrees of change among the “target“ groups. The lower the degree of change in species composition between plots in the “target“ groups relative to that of vascular plants, the higher the percentage “target“ species encompassed in a complementary selection of sites based on vascular plants. We conclude that in practical site selection of small scale sites of conservation value, such as woodland key habitats, vascular plants may be used in combination with an inventory of important habitats for rare and/or redlisted forest species, such as dead wood, old trees, deciduous trees, and cliffs.

Sammendrag

In this study of 130 sites with different management we investigated whether vascular plant species richness is significantly reduced when traditionally managed hay meadows are abandoned and reforested. We also compared the effects of reforestation with those of intensified land-use to see which have the largest effects on species richness. Finally, we investigated the relative importance of relevant ecological factors for species richness.While the use of artificial fertilizers in traditionally managed hay meadows has resulted in significantly lower species richness, and intensive cultivation in even lower species richness, abandonment with reforestation has not decreased the species richness significantly.Productivity and habitat diversity have determined the species richness of meadows on the scale (0.035.1 ha) of this study. Low productivity is a prerequisite for high species richness in meadows. Maximum species richness was observed in unproductive, old, traditionally managed hay meadows with a high soil pH and high habitat diversity. The high species richness of these meadows suggests that they are in urgent need of conservation.

Sammendrag

Epiphytic lichens (and some non-lichenized fungi) on 34 coppices (204 stems) of Corylus avellana were investigated in a 140 ha study area in south-western Norway. A total of 65 species were recorded on a total bark area of 63 m2. Corylus in broad-leaved deciduous forest supported more species of macrolichens, and fewer species of icrolichens, than Corylus in pine forest. The macrolichen flora of the deciduous forest differed from that of the pine forest by having a rich flora of species belonging to the Lobarion alliance. Old Corylus coppices with tall stems (>8 m), large girth (>8 cm diameter at breast height) and a noticeable cover of macrolichens (>10% of bark area) supported the highest number of rare species, and overall, species of mcrolichens. More than 50% cover of icrolichens indicated richness and rarity of microlichens on Corylus

Sammendrag

We compared diversity of birds in 35 study plots of equal size (58 ha) and productivity in western Norway, ranging from pure native pine Pinus sylvestris forests (n = 7), through different mosaics of native pine forests and spruce Picea spp. plantations (n = 21), to pure spruce plantations (n = 7). Diversity was evaluated by means of species richness, diversity indices, relative abundance curves and rarefaction. The diversity indices appeared to be less suitable for our purpose. Species richness was higher in pine forest than in spruce forest. However, a peak in species richness was found in mosaic forest. For pooled samples (408 ha), 11 bird species recorded in pine forest were not found in spruce forest, seven species were found in spruce forest but not in pine forest, and seven species were confined to the medium mosaics of pine and spruce forest (on average 56% pine and 44% spruce). We argue that, when mixing two habitat types A and B, the ratio of these habitats that maximize avian diversity depends on the ratio of species confined to habitat A and B, as well as the number of species favoured by the mixture of A and B. Existing spruce plantations (13% of the area) in native pine forests of western Norway have reduced the diversity of birds locally, but increased the diversity of birds on the landscape and regional scale.