Biodiversity and Conservation 11: 1 479 1503 , 2002. 2002 Kluwer Academic Publishers. Printed in the Netherlands.

Habitat preferences of red-listed fungi 1 and bryophytes in woodland key habitats in southern Sweden analyses of data from a national survey
3 3 AKE BERG 1,2, *, ULF GARDENFORS , TOMAS HALLINGBACK and 4 MIKAEL NOREN

Department of Conservation Biology, SLU, Box 7002, S-750 07 Uppsala, Sweden; 2 The Swedish Biodiversity Centre, SLU, Box 7007, S-750 07 Uppsala, Sweden; 3 Swedish Threatened Species Unit, 4 , Sweden; SLU, Box 7007, S-750 07 Uppsala, Sweden; National Board of Forestry, S-551 83 Jonkoping * Author for correspondence (e-mail: ake.berg@ nvb.slu.se; fax: 146 -18 -673537) Received 20 February 2001; accepted in revised form 30 August 2001

Key words: Bryophytes, Forest stand composition, Fungi, Geographical location, Ground conditions, Historical land-use, Lichens, pCCA, Woody substrates Abstract. The aim of this study was to identify habitat preferences of red-listed epiphytic and epixylic bryophyte, lichenized and non-lichenized fungi species in woodland key habitats (WKHS) (areas less than 10 ha, where forest structures indicate occurrence of red-listed species) in southern Sweden. The relative importance of different groups of environmental factors was assessed with partial canonical correspondence analysis techniques and a cross-validation approach using data from 7196 selected WKHs. Different woody substrates (old trees, logs and snags) made up the most important variable group for occurrence of red-listed species (30% unique explainable variation). Species associated with Fagus sylvatica and Picea abies habitats, but also species associated with Quercus spp. and Populus tremula habitats showed distinct habitat preferences. The second most important variable group (16% unique explainable variation) was geographical location. A westeast gradient was identied, and species concentrated to Baltic islands in the east were separated from other species. This gradient, and an identied southnorth gradient, probably reect differences in temperatures and rainfall between different regions. Among the remaining variable groups, historical land-use, ground conditions and forest stand composition were of similar importance (57% unique explainable variation). Traditional management regimes resulting in semi-open forest habitats (leaf harvesting, forest grazing and selective cutting) were associated with the occurrence of many species, probably due to differences in microclimate between sites of different openness. Furthermore, a ground moisture gradient extending from species associated with dry sites (mainly lichenized fungi) to species associated with wet sites (mainly bryophytes), and a nutrient gradient from species associated with nutrient-poor sites to species occurring at nutrient-rich sites, were identied. Thus, conservation measures are needed in a broad spectrum of habitats with different substrates. Also sites with similar substrates, but situated in different regions (and climates), or with different ground moisture and nutrient conditions are needed to cover the full spectrum of habitat conditions suitable for different red-listed bryophytes and fungi.

Introduction Swedish forests have been used by forestry for centuries, and especially the forests
1

Includes both lichenized fungi (lichens) and non-lichenized fungi.

1480 in southern and central Sweden have been heavily inuenced by forestry operations (Tenow 1974; Bernes 1994; Hansson 1997). Modern forestry with clearcutting, cleaning and thinning started during the 1950s and has changed the forest landscape drastically (Esseen et al. 1992), and today only 3% of the productive forestland has escaped intensive harvesting (Ostlund et al. 1997). As a consequence, plant and animal communities have changed considerably and many threatened species are restricted to remnants of natural forests or substrates typical of old forests. Earlier studies of threatened species in Swedish forests have suggested that lack of old trees and dead wood, low proportion of deciduous trees (Berg et al. 1994, 1995; Rydin et al. 1997; Jonsell et al. 1998) and restricted areas of old forests on soils with high fertility (Gustafsson 1994; Rydin et al. 1997) are major causes of the decline and rarity of many populations of forest plants and animals. Modern forestry, with short rotation periods and planting of monocultures of coniferous trees, has been suggested to be a major cause of the decline of many red-listed species (e.g. Berg et al. 1994, 1995), i.e. species in risk of extinction. Swedish forestry policy changed markedly in 1993 when the earlier production goal and a new environmental goal were given equal importance, whereby biodiversity was to be secured and ecosystems conserved. These new goals were mainly to be achieved by the forestry sector itself. Therefore, a countrywide survey of woodland key habitats (WKHs) on privately owned forestland (1.2 3 10 5 km 2 or half of all forestland in Sweden) was initiated in Sweden in 1993 (Nitare and Noren 1992) by the government and was completed in 1998. A WKH is dened as an area (mostly less than 10 ha) where one or more nationally red-listed species occur, or where forest structures indicate a strong likelihood for such an occurrence (Nitare 1992). Red-listed species in Sweden encompass threatened species as and Noren well as a category of lower threat denoted Near Threatened (earlier termed Care Demanding), including ca. one-third of the red-listed species (Aronsson et al. 1995; Gardenfors 2000). The main task of the WKH survey was to localise, delimit and describe WKHs on all privately owned forestland. During the survey, 40 071 WKHs with a total area of 1187 km 2 were investigated, which makes it one of the largest nature conservation surveys in the world (National Board of Forestry 1998). A later validation survey revealed that a majority of the sites (71%) harboured red-listed bryophyte or lichenized fungi species (Gustafsson et al. 1999). On the other hand, red-listed vascular plants seemed to be as rare in WKHs as in surrounding production forests (Gustafsson 1999). However, identied WKHs seem to fulll their initial goal, i.e., to harbour red-listed species. Species with habitat selection on a larger landscape scale (e.g. many mammals and birds) could not, on the other hand, be expected to be restricted to small habitat fragments such as the studied WKHs. A large number of observations (a magnitude of 10 5 observations) of red-listed species (according to the classication in Aronsson et al. 1995), mainly bryophytes, lichenized and non-lichenized fungi, have been made (National Board of Forestry 1998). This offers unique possibilities to analyse patterns of occurrence, quantify the importance of different environmental factors, and to identify conservation measures and priorities for Swedish forest landscapes. Earlier studies of habitat

1481 requirements of red-listed species have largely been based on opinions of experts on different taxa (see e.g. Berg et al. 1994, 1995; Gustafsson 1994; Berg and Tjernberg 1996; Rydin et al. 1997; Jonsell et al. 1998), or species-by-species presentations of habitat associations and threats to red-listed species (Larsson 1997; Hallingback 1998; Thor and Arvidsson 1999). The aim of the present study was to identify broad-scale habitat preferences of easily identied red-listed epiphytic and epixylic bryophytes, lichenized and nonlichenized fungi in WKHs in southern Sweden. The relative importance of different variable groups (geographical location, stand composition, forest structure, woody substrates, ground conditions and present and historical land-use) was assessed with ordination techniques and a cross-validation approach. Based on the present and similar studies, a desirable goal in forest conservation efforts is to be able to predict occurrences of cryptic or hard to identify red-listed species from easily available environmental data. Implications of the ndings for forest management and conservation are discussed, and future investigations in WKHs are suggested.

Methods The survey of woodland key habitats The main aim of the WKH survey was to identify and describe the WKHs. Two main criteria for identication of WKHs were occurrence of (1) structural elements, such as old trees, logs and snags a priori classied as suitable for red-listed species, and (2) presence of indicator species (including not red-listed species), mainly et bryophytes, fungi and vascular plants with habitat demands at stand level (Noren al. 1995). The personnel doing the eldwork were qualied silviculturists who had been especially trained in the identication of threatened habitats and red-listed species. Halfway through the survey an evaluating assessment was made in order to control and secure uniform quality of the eldwork. Complete censuses of different taxa were not made, but all observations of red-listed species and indicator species [easily identied, relatively common species indicating high conservation values (see National Board of Forestry 1998)] were noted. Large numbers of observations of red-listed species were made and red-listed species were observed in 41% of the WKHs in southern Sweden (National Board of Forestry 1998). A large proportion of these were bryophytes, lichenized and non-lichenized fungi (ca. 95%) and therefore sites with observations of red-listed bryophytes and fungi were selected for the present study. We chose to delimit the study to southern Sweden (see Figure 1). One important reason for this delimitation is that forest companies and the Swedish state, whose censuses were not included in the WKH survey, own large tracts of forest (ca. 60%) in northern Sweden (Statistics Sweden 1999). Furthermore, large continuous areas of WKHs, and occurrence of different habitat types (e.g. mountain forests) in northern Sweden made it more difcult to apply standardised WKH census methods there.

1482

Figure 1. Map of the counties of southern Sweden that were included in the study (dark grey). The dark line delimits the two major regions (Gotaland and Svealand) in southern Sweden.

Databases and analyses Initially, nearly 8600 WKHs with observations of red-listed bryophytes, lichenized and non-lichenized fungi, located on privately owned small-scale forestry areas in southern Sweden, were selected from databases (a total of ca. 21 000 WKHs in southern Sweden) at the National Board of Forestry. This reduction of the database was performed in order to delimit the study to sites that were shown to harbour red-listed species at present. Further reductions of the database due to lack of data for the selected habitat variables (see below), as well as exclusion of sites where only very rare red-listed species (found at , 30 sites) were observed, resulted in a nal set of 7196 WKHs. The original databases included over 250 habitat variables (see National Board of Forestry 1998). However, many of these measured similar attributes and others were considered to be of restricted interest for the present study. Therefore, many variables were combined or omitted, and a nal set of 130 habitat variables was selected for exploratory analyses (Table 1). These variables were grouped into six main categories (geographical location, stand composition, forest structure, woody substrates, ground conditions, and present and historical land-use). The restricted censuses do not make usage of standard statistical methods feasible, since non-occurrences in reality could be occurrences. However, the canonical correspondence analyses (CCAs) emphasise occurrences and this tech-

1483
Table 1. Description of habitat variables in the six main variable groups. Variables in parentheses were excluded from further analyses due to correlations with other habitat variables or weak associations to the identied axes in the exploratory analyses. Habitat variables and variable categories Description

Geographic location (including transformations) Longitude (X ) Swedish national grid coordinates for Latitude (Y ) the centre of each site with 10 m precision X 2, X 3, Y 2, Y 3, X 2Y, XY 2 Square and cubic transformations Woody substrates Abundance of logs of: Populus, Picea, Pinus (all coniferous trees), all other deciduous trees a and all southern deciduous trees b (Snags) (Tall stumps) Woody debris (Standing dead trees) Number of old trees of: Ulmus glabra, Ul. minor, Fraxinus, Populus, Fagus, Corylus, Tilia, Acer, Quercus, (Pinus), (all coniferous trees) and all deciduous trees Trees with wood mould (decayed inner parts of the stem) Slow growing Quercus (Slow growing trees of all species) Large trees of: Ulmus, (Fraxinus), (Populus), (Fagus), ( Quercus), (Picea), (Alnus), (Acer), Tilia, (Salix), (Pinus), Betula, (all deciduous trees) Historical and present land-use Earlier selectively cut forest Earlier forest grazing Earlier leaf harvesting Undisturbed natural forest (Earlier mowing for hay) (Earlier clearcutting) Present grazing Occurrence of pollarded trees (Fire refugia) Forest re area Ground conditions Spring Moisture Heterogeneity in moisture Proportions of different ground vegetation types: Lichen dominated, (lichen rich), (poor dwarfshrub dominated), (Empetrum Calluna dominated), (Vaccinium vitis idaea dominated), Vaccinium myrtillus dominated, Equisetum -Carex dominated, dominated by broadleaf grasses, dominated by narrowleaf grasses, bare ground dominated, low herb dominated, tall herb dominated and rich herb dominated

Ordinal scale 13 Ordinal scale 13

Ordinal Ordinal Ordinal Ordinal

scale scale scale scale

13 13 13 13

Ordinal scale 13

Presence / absence Presence / absence Presence / absence Number of large trees above species specic size limit (National Board of Forestry 1998)

Presence / absence Presence / absence Presence / absence Presence / absence Presence / absence Presence / absence Presence / absence Presence / absence Presence / absence Presence / absence

Presence / absence Ordinal scale 14 of dominating moisture class Difference in moisture classes within a site (03) Ordinal scale (10% cover of different ground vegetation types)

1484
Table 1. (continued ) Habitat variables and variable categories Boulder-rich ground Outcrop of bedrock Slopes (Moving groundwater) (Flooded forest) Calcareous ground Forest structure and composition Tree volume (of all species) Tree density Forest stratication Forest age Proportion of different tree species: Fraxinus, Populus, (Carpinus), Fagus, Quercus, Betula, Picea, Corylus, Alnus, Acer, (Tilia), (Pinus), (Ul. glabra) Ul. minor, and (all deciduous trees combined) (Tree species diversity) Stand composition WKH area Proportion of main habitats: Production forest, (mire), rocks, patches of arable land within forest, (water) and (other habitats)
a

Description Presence / absence Presence / absence Presence / absence Presence / absence Presence / absence Presence / absence Ordinal scale (m 3 forest in 50 m 3 intervals / ha) Ordinal scale 14 Stratication or not of tree strata Age class (10 years) of dominating trees Ordinal scale (10% cover classes) of different dominating tree species

Simpson index Area (ha) Proportion of main habitats (10% classes)

All other deciduous trees includes deciduous trees found over most of Sweden (Populus tremula, Betula spp., Alnus glutinosa, Sorbus spp., Salix spp. etc.). b All southern deciduous trees includes species restricted to southern Sweden (Fagus sylvatica, Quercus robur, Quercus petraea, Ulmus spp., Tilia cordata, Fraxinus excelsior, Acer platanoides etc.).

nique has earlier been used on incomplete species data (Hallgren et al. 1999). Results from the latter study and exploratory analyses of the WKH data set (see below) suggest that meaningful evaluations of species in relation to habitat gradients can be made using such data. However, evaluations of site scores are not meaningful, since complete species data does not exist for all sites. Noisy environmental data have been shown to have a strong inuence on CCA, i.e., non-existing gradients are easily identied (McCune 1997). Therefore, we used CCAs on a divided data set with explanatory analyses and hypothesis formation (by using an explanatory data set) and hypothesis testing (by using a conrmatory data set, cf. Hallgren et al. 1999). The nal evaluation of gradients was made on the full database (i.e. both halves) in order to include as much information as possible in the analyses (as suggested for predictive models by Fielding and Bell 1997). The analyses were conducted with CANOCO 4 (ter Braak and Smilauer 1998). However, limitations set by the software made it impossible to use all 7196 sites in a few of the nal partial canonical correspondence analyses (pCCA) on the full database, and a maximum of 2300 sites had to be randomly excluded in a few analyses. Stepwise CCA (ter Braak 1987; ter Braak and Smilauer 1998), which selects the most signicant variables one at a time, was used to identify important habitat variables and the order in which the main variable groups entered the models. CCA

1485 was used to investigate if one group of variables could explain any variation in species composition not explained by another set of habitat variables (Legendre and Legendre 1998; ter Braak and Smilauer 1998). The different categories of habitat variables (Table 1) to some degree contain the same information, e.g. geographical location and tree species composition (i.e. different species dominate in different parts of southern Sweden). In order to quantify the unique contribution of each class of variables in explaining species composition, we performed variation partitioning on the models for the entire data set (kland and Eilertsen 1994; Roche et al. 1998). We partitioned the variation explained by the explanatory variables, since total inertia has been suggested to be an inadequate measure of the total variation in the data set (kland 1999). However, only selected combinations of variable groups that were of special interest were included in the variation partitioning due to the relatively large number of variable groups. In the initial CCAs we found that species scores were most easily interpreted when the presence / absence of species was used, and therefore available abundance estimates of different species were not used. Furthermore, species scores and environmental axes were most easily interpreted when variables that were recorded as proportions (e.g. abundance of tree species, ground vegetation types) were arcsine transformed (Legendre and Legendre 1998). The exploratory analyses also showed that two variables (logs of coniferous trees and old coniferous trees) were highly correlated with other habitat variables and these variables were therefore omitted from the analyses. Furthermore, a relatively large number of variables (33) that were weakly associated with the identied axes (t -value , 2.1) could not be expected to contribute to the t of the species data (ter Braak and Smilauer 1998) and were therefore excluded from the following analyses (excluded variables, see Table 1). Still, a large number (70) of habitat variables remained, but the number of sites was much larger (limitations in variable numbers, see ter Braak and Smilauer 1998), and associations with variables were mainly related to categories of several habitat variables (Table 1).

Results Exploratory analyses In order to identify the importance of the different groups of habitat variables, we performed a series of stepwise CCAs. Generally, several variables from the same variable category entered the models before those from other categories. The variable groups entered the models in the following order: woody substrates, geographical location, historical and present land-use, ground conditions, forest composition, and nally stand composition. We therefore performed a series of sequential analyses where the importance of these categories was tested in sequence after removing effects of other variable groups regarded as more important (Table 2). These analyses suggested that the occurrence of bryophytes, lichenized and

1486 non-lichenized fungi in the censused WKHs was strongly associated with the occurrence of suitable woody substrates and geographical location. There were also substantial effects of ground conditions, forest structure, and present and historical land-use, while the effect of stand composition was weak. These hypotheses (Table 2) were subsequently tested on the conrmatory data set (see below). Conrmatory analyses The proposed hypotheses were tested with Monte Carlo simulations (199 permutations) of all canonical axes using Holms adjustment (Legendre and Legendre 1998) for multiple comparisons on the conrmatory data set. All statistical tests resulted in acceptance of the proposed hypotheses (see Table 2), so we conclude that the included habitat categories all were related to the occurrence of the studied redlisted species in WKHs in southern Sweden. Combined analyses based on the entire database Occurrence of different woody substrates was the most important group of factors for occurrence of the studied epixylic and epiphytic species. The rst axis of the
Table 2. Hypotheses and results from statistical tests on all canonical axes on the conrmatory data set. The sequential analyses and hypotheses had been proposed by the exploratory analyses. Holms adjustments consider six comparisons (see Legendre and Legendre 1998). Hypothesis Environmental variables Woody substrates Covariables Calculated P -value 0.005 Holms adjusted P -value 0.03

Red-listed species are associated with amount of different woody substrates Red-listed species are associated with latitude and longitude Red-listed species are associated with historical and present land-use Red-listed species are associated with ground conditions Red-listed species are associated with forest stand composition Red-listed species are associated with stand composition

Latitude and longitude

Woody substrates

0.005

0.03

Land-use

Latitude and longitude Woody substrates Latitude and longitude Woody substrates Land-use Latitude and longitude Woody substrates Land-use Ground conditions Latitude and longitude Woody substrates Land-use Ground conditions Forest composition

0.005

0.03

Ground conditions

0.005

0.03

Forest composition

0.005

0.03

Stand composition

0.005

0.03

1487 CCAs, with abundance of different woody substrates as environmental variables, consisted of a gradient from species associated with old oaks Quercus robur / petraea (including many lichenized fungi, e.g., Cliostomum corrugatum, Ramalina baltica, and Sclerophora coniophaea) and beech Fagus sylvatica (e.g. the bryophyte Neckera pumila and the lichen Biatora sphaeroides) to species associated with Norway spruce Picea abies and aspen Populus tremula, which mainly included bryophytes and non-lichenized fungi (Figure 2). The groups associated with aspen and spruce were also separated along the rst ordination axis, i.e., species associated with spruce (e.g. the bryophyte Lophozia ascendens and the fungus Phellinus nigrolimitatus) were found together in the biplot, while species associated with aspen logs (e.g. Clavicorona pyxidata and Antrodia pulvinascens) and old aspen (e.g. P. populicola and N. pennata) were found in the lower right quartile of the biplot (Figure 2). Species associated with beech were clearly separated from the other species along the second axis (Figure 2). Species associated with other deciduous trees and Scots pine Pinus sylvestris were found at the centre of the biplot, suggesting that these species were not so strongly associated with specic woody substrates. Geographical location was the most important group of factors for occurrence of different red-listed species after covarying out occurrence of different woody substrates (i.e. using woody substrates as covariables in pCCA). The rst axis of the pCCA with geographical coordinates as environmental variables was interpreted as a westeast gradient and the second axis as a southnorth gradient (Figure 3). Most northern species (e.g. Bryoria bicolor, Collema subnigrescens and Chaenotheca gracillima) occurred together in the upper left quartile of the biplot. Eastern species (e.g. Dicranum tauricum, Inonotus hispidus and Gyalecta truncigena) occurred at the right part of the biplot, while south-western species (e.g. Holwaya mucida, Normandina pulchella and Opegrapha vermicellifera) occurred in the lower left part of the biplot (Figure 3). There were no apparent differences in the distribution patterns of the three main taxa (bryophytes, lichenized and non-lichenized fungi). Historical land-use was classied as the most important factor for occurrence of different red-listed species, after covarying out geographical location and occurrence of different woody substrates. The rst axis of the pCCA with different forms of land-use as environmental variables separated species associated with leaf harvesting and occurrence of pollarded trees (mainly lichenized fungi, e.g., S. farinacea, G. ulmi and Biatorella monasteriensis) from the other species (Figure 4). The second ordination axis seemed to consist of a gradient extending from relatively open sites (at the top of the biplot), including the mentioned species associated with leaf harvesting, but also species associated with earlier-grazed forests or sites used for selective cutting (e.g. Microcalicium ahlneri, Catillaria sphaeroides and Antrodia pulvinascens) to species found in dense forests (e.g. Pertusaria multipuncta, Dicranodontium denudatum and O. viridis) at the bottom of the biplot (Figure 4). In general, several lichenized and non-lichenized fungi species were associated with sites having these types of old management regimes, while most bryophytes were found at sites where these management regimes have not been practised. Ground conditions were classied as the most important variable group for

1488

Figure 2. Ordination biplot (CCA) with woody substrates as environmental variables. The locations of species have in some cases been shifted slightly for clarity. Slowoak abundance of slow-growing oaks. Five environmental variables with weak associations to the identied axes are excluded from the biplot. 7SPEC indicates approximate position of Pac tube, Bue viol, Ope ille, Cha pae, Cli corr, Fis hepa and Cal luci. 4SPEC indicates position for Eve diva, Mic ahln, Bry bico and Cal parv. For full species names see Appendix.

1489

Figure 3. Ordination biplot (pCCA) with geographical location as environmental variables and woody substrates as covariables. The locations of species have in some cases been shifted slightly for clarity. 7SPEC indicates approximate position of Ant pulv, Phe ferf, Ram balt, Cla pyxi, Cal quer, Scli coni and Cli corr. For full species names see Appendix. For explanation of environmental variables, see Table 1.

1490

Figure 4. Ordination biplot (pCCA) with historical land-use as environmental variables and woody substrates and geographical location as covariables. PollTrees occurrence of pollarded trees, Leafharvest signs of earlier leaf harvesting, Grazing earlier grazed forests, Undisturbed not cut forest and SelectCut selectively cut forest. The locations of species have in some cases been shifted slightly for clarity. 6SPEC position for Pyr nita, Bue viol, Fis hepa Sch peri, Cli corr and Phe ferf. 3SPEC position for Cla pyxi, Ram balt and pac tube. 3 SPECIES position for Phe nigl, Lca glab and Scl coni. For full species names see Appendix.

1491 occurrence of different red-listed species when using coordinates, woody substrates and land-use as covariables in pCCA. The rst axis of the pCCA with ground condition as environmental variable consisted of a moisture gradient extending from species associated with relatively dry sites (e.g. Cladonia parasitica, Evernia divaricata and B. bicolor) to species associated with moist or wet sites (e.g. Trichocolea tomentella, Menegazzia terebrata and Arthonia spadicea). In general, bryophytes were associated with moist sites, while lichenized fungi were associated with relatively dry sites (Figure 5). The second axis was interpreted as a nutrient gradient from species associated with nutrient-poor sites (e.g. M. terebrata and C. gracillima) to species occurring at nutrient-rich sites (e.g. T. tomentella, S. peronella and Skeletocutis nivea), see Figure 5. Forest stand composition was classied as the most important factor for occurrence of different red-listed species when using coordinates, woody substrates, land-use and ground conditions as covariables in pCCA. The rst axis of the pCCA with forest stand composition as environmental variable consisted of a forest age gradient, extending from species associated with moderately old forests (ca. 100 years) with a high volume of trees (e.g. Usnea orida, Sk. nivea and B. bicolor) to species associated with very old forests (ca. 180 years) with a smaller volume of trees (e.g. Cl. corrugatum, C. phaeocephala and R. baltica), see Figure 6. In general, lichenized fungi were associated with older forests than bryophytes and nonlichenized fungi. The second axis was difcult to interpret. Stand composition was signicantly associated with the occurrence of different red-listed species when using all other variable groups as covariables in pCCA. The rst axis of the pCCA with stand composition as environmental variable consisted of a topographic gradient, from species associated with occurrence of rocks (including both species using rocks as a substrate and species associated with trees in rocky areas, e.g. B. bicolor, O. illecebrosa and Cla. parasitica) to species occurring in plain forests (e.g. Bi. sphaeroides, U. orida and Sk. nivea), see Figure 7. The second axis consisted of a gradient extending from species associated with sites with patches of arable land within forests (e.g. G. truncigena, M. terebrata and Inonotus hispidus) to species associated with continuous forest (E. divaricata and Bia. monasteriensis). There were no apparent differences in the effects of stand composition on the three main taxa. Variation partitioning Variation partitioning was only performed for certain combinations of variable groups due to the relatively large number of groups. First, the unique variation explained by each variable group (i.e. when effects of all other variable groups were removed) was calculated for all six variable groups. These calculations suggested that woody substrate variables accounted for the largest amount of unique explainable variation (30.2%). Geographical location also accounted for a large proportion of unique variation (15.8%), while the unique variation explained by ground conditions (6.6%), stand composition (6.0%) and land-use (4.8%) was smaller. The variation accounted for by stand composition (1.1%) was small. Thus, 64.5% of the

1492

Figure 5. Ordination biplot (pCCA) with ground conditions as environmental variables and woody substrates, geographical location and historical land-use as covariables. Vshortherb proportion with short herb vegetation, Vtallherb proportion with tall herb vegetation, VrichHerb proportion with rich herb vegetation,Vvaccin proportion with Vaccinium myrtillus dominated vegetation and VcarexEquis proportion with Carex or Equisetum dominated vegetation. The locations of species have in some cases been shifted slightly for clarity. 4SPEC position for Tric tom, Dicr den, Buxb vir and Ope ochr. For full species names see Appendix.

1493

Figure 6. Ordination biplot (pCCA) with forest composition as environmental variable and woody substrates, geographical location, historical land-use and ground conditions as covariables. The locations of species have in some cases been shifted slightly for clarity. 4 SPEC Ope viri, Phe fere, Pyr nita and Lca glab. For full species names see Appendix.

1494

Figure 7. Ordination biplot (pCCA) with stand composition as environmental variable and woody substrates, geographical location, historical land-use, ground conditions and forest composition as covariables. The locations of species have in some cases been shifted slightly for clarity. 4SPEC position for Per mult, Cha phae, Ram balt and Cal quer. 3SPEC position for Ope sore, Anas hel and Buxb vir. For full species names see Appendix.

1495
Table 3. Partitioning of the explained variation among the four different habitat composition variable groups (woody substrates, ground conditions, forest composition and stand composition)a . A B Percentage of explained variation AuB Woody substrates Ground conditions Forest composition Stand composition Ground conditions, forest composition Ground conditions, stand composition Forest composition, stand composition Ground conditions, forest composition, stand composition Forest composition Stand composition Stand composition, forest composition Forest composition, woody substrates Woody substrates, stand composition Woody substrates, stand composition, Forest composition Stand composition Woody substrates, ground conditions Woody substrates, stand composition Ground conditions, stand composition Woody substrates, ground conditions, Stand composition Woody substrates, ground conditions Woody substrates, forest composition Ground conditions, forest composition Woody substrates, ground conditions, forest composition 60.7 50.8 72.3 48.9 60.3 50.6 48.4 11.8 24.4 11.7 9.6 12.4 9.5 37.2 12.9 15.5 24.6 12.7 1.8 1.7 1.9 1.6 A>B 12.4 22.4 0.9 24.3 12.9 22.6 24.8 12.9 0.4 13.0 15.1 12.4 15.2 0.7 25.1 22.4 13.3 25.2 0.8 0.9 0.8 1.0 BuA 12.3 15.6 1.8 25.4 14.1 17.2 26.8 25.0 2.3 26.9 73.6 62.8 75.3 1.9 60.4 52.5 13.7 62.1 84.7 87.8 49.0 97.4 A<B 85.5 88.8 74.9 98.6 87.3 90.4 100.0 49.8 27.0 51.6 98.4 87.6 100.0 39.9 98.4 90.4 51.6 100.0 87.3 90.5 51.6 100.0 Other 14.5 11.2 25.1 1.4 12.7 9.6 50.2 73.0 48.4 1.6 12.4 60.1 1.6 9.6 48.4 12.7 9.5 48.4

Ground conditions

Forest composition

Stand composition

AuB is the variation explained by A but not by B. A>B is the variation jointly described by A and B. BuA is the variation explained by B but not by A. A<B is the variation described by A and B together.

explainable variation was unique for different variable groups, while the remaining 35.5% of this variation was explained by several of these variable groups together. Variation partitioning of the four habitat variable groups measuring habitat composition (woody substrates, ground conditions, forest composition and stand composition) suggests that the unique variation explained by these variables together was 52%. A complete variation partitioning for the four habitat composition variable groups is presented in Table 3.

Discussion The occurrence of high quality substrates such as old living trees and logs is of vital importance for most red-listed bryophytes and fungi in the censused WKHs (30% unique explainable variation). This is in concordance with the results from earlier

1496 compilations of expert knowledge (e.g. Berg et al. 1994, 1995; Rydin et al. 1997), suggesting that occurrence of suitable substrates is the major factor restricting the occurrence of red-listed Swedish forest species. It must, however, be stressed that some taxa (e.g. mycorrhizal fungi) with different habitat preferences than the studied species were not included in our study. The present study suggests that red-listed species associated with beech, oak, spruce or aspen substrates have distinct habitat preferences and rarely occur at sites dominated by other substrates. Species associated with other tree species were not strongly associated with any single tree species, but other factors (e.g. earlier land-use and soil conditions) were more important. Thus, ve major WKH types (dominated by beech, oak, spruce, aspen and a group with the remaining tree species, respectively) with complementary ora of red-listed species could be identied, although the transition between some groups is gradual. Of these WKHs, habitats dominated by beech, oak or other broad-leaved deciduous trees harboured the largest number of red-listed species, while fewer species were associated with spruce- or aspen-dominated habitats (Figure 2). The second most important factor (16% unique explainable variation) for the occurrence of red-listed bryophytes and fungi in the studied WKHs was geographical location, which partly consisted of a westeast gradient (Figure 3). An analysis of the total number of red-listed species in different regions of Sweden (data from Gardenfors 2000) reveals that the eastern islands Oland and Gotland also have very large numbers of red-listed species unique for each province [and also large numbers of red-listed species in total (including species occurring in other regions)], in relation to their small areas. Similar results have also been found for vascular and Borgegard 1999). plants (Rosen Geographical location also included a southnorth gradient. There are considerable climatological differences between different regions in southern Sweden (e.g. precipitation, mean temperature), which result in differences of up to 30 days in the length of the growing season, but also in climatological differences between coastal areas and continental interior areas (Raab and Vedin 1995). Furthermore, the southnorth gradient seems to reect differences in tree species composition between regions. For instance, many of the species restricted to southernmost Sweden (Skane, Halland and Blekinge) were associated with different southern deciduous trees that are much more abundant in these regions than in central Sweden (Statistics Sweden 1999). Similarly, species associated with aspen were more abundant in central than in southernmost Sweden, probably because aspen is twice as abundant in the boreonemoral zone (a mean of 2.3% of the tree volume, but up to 6% of the tree volume in some regions around Lake Malaren) as in the nemoral zone (1.2% of the tree volume, see Statistics Sweden 1999). However, the distribution of several species associated with spruce and oak did not reect the present abundance of these trees in different regions. Species associated with spruce were more common in central Sweden than in southern Sweden, despite the fact that spruce is as common in the nemoral as in the boreonemoral zone (ca. 50% of the tree volume (see Statistics Sweden 1999)). However, spruce has recently colonised (and

1497 been planted in) southernmost Sweden (Berglund et al. 1996) and a probable reason for the restricted occurrence of red-listed species associated with spruce in these regions is the lack of old forests with continuous supply of suitable spruce substrates. Similarly, species associated with oak were more abundant in central than in southernmost Sweden, despite the fact that oak trees were more abundant in southernmost (ca. 3.2% of the tree volume) than in central Sweden (ca. 1.5% of the tree volume (see Statistics Sweden 1999). Possible reasons are the higher levels of air pollution in southernmost Sweden (e.g. Thor 1998), differences in stand composition or differences in continuity of occurrence of suitable substrates between regions. Historical land-use, ground conditions and forest composition were all of similar importance (57% unique explainable variation) for the bryophyte and fungi ora. First, species associated with leaf harvesting were separated from the other species. The pollarded trees with red-listed species were mainly trees such as ash Fraxinus excelsior, lime Tilia cordata and elm Ulmus glabra. Historically, leaf harvesting was a widespread agricultural practice throughout most of Sweden (Slotte 2000). Today, trees that earlier were pollarded, often situated in landscapes that no longer are managed in a traditional way, are refuges for several threatened species (see also Moe and Botnen 1997). Historically, these species were found in extensive areas of natural meadows and pastures that are now overgrown, planted with forest or used as arable elds (Slotte 2000). Pollarded trees are often very old, with decaying wood and cavities, and offer a complex substrate with large variation in microclimate and nutrient conditions within the same tree. Such trees are of great conservation value also for taxa such as insects (Berg et al. 1994). Leaf harvesting of deciduous trees, and grazing or mowing of natural grasslands in semi-open agricultural landscapes, is therefore of great conservation value also for forest species and must be promoted by subsidies since there is no economic incitement for such a practice. Additionally, associations with historical land-use consisted of a gradient extending from relatively open sites, including the species associated with leaf harvesting as well as species associated with earlier grazed forests or selectively cut sites, to species in closed forests (Figure 4). Generally, most non-lichenized fungi were associated with grazed or cut forests, while most bryophytes were found in dense, earlier not managed forests. These differences between taxa probably depend on biological differences regarding preferences for climatic factors, such as humidity and sun exposure, that differ between open and dense forests. Earlier studies have also suggested that threatened macrofungi are often found in mesic to dry habitats (Rydin et al. 1997), whereas lichenized fungi in general have been found to be most frequent in open dry sites (Gustafsson and Eriksson 1995). Many red-listed invertebrates in Sweden have also been shown to prefer open forest and trees in exposed positions, for instance species associated with oak, while species associated with beech frequently prefer shaded forests (Gardenfors and Baranowski 1992; Jonsell et al. 1998). Most bryophytes, on the other hand, are favoured by humid conditions found in dense or protected sites (see also Gustafsson et al. 1992; Gustafsson and Eriksson 1995). Thus, the openness of the habitat and the microcli-

1498 mate is important for many forest organisms. Management regimes resulting in semi-open forest habitats (leaf harvesting, forest grazing and selective cutting) are not used in modern forestry and farming. Among ground condition variables there was a moisture gradient from species associated with relatively dry sites to species associated with moist or wet sites (Figure 5). Again, bryophytes were mainly associated with moist sites, while lichenized fungi were associated with relatively dry sites. Furthermore, there was a nutrient gradient extending from species associated with nutrient-poor sites to species occurring at nutrient-rich sites. This suggests that there is an effect of nutrient conditions on the occurrence of red-listed bryophytes, lichenized and non-lichenized fungi which is not associated with occurrence of suitable substrates or large regional differences. Most of the studied species are found on dead wood or old trees, suggesting that bark chemistry is affected by nutrient conditions in the ground, possibly by transport of nutrients from the roots to the bark. This has been suggested in a study of the epiphytic ora on aspens (Gustafsson and Eriksson 1995), in which eld vegetation type correlated strongly with chemical properties of the bark and in the ground. Thus, sites with similar substrate types, but on different soils, harbour different rare bryophytes, lichenized and non-lichenized fungi, e.g. Lobaria pulmonaria and U. longissima have been shown to be associated with bark of trees situated in different soil conditions (Gauslaa 1995; Gauslaa et al. 1998). Protection and management of a spectrum of sites with similar substrates but different soil conditions are therefore needed. Among forest composition variables there was a forest age gradient, extending from species associated with forests aged 100 years to species associated with forests aged 180 years (Figure 6). However, all WKHs consist of more or less old forests (or forests with some old trees) compared with Swedish forests in general (88% of the forests in southern Sweden are younger than 100 years; see Statistics Sweden 1999). When comparing organism groups, lichenized fungi were more associated with older forests than bryophytes and non-lichenized fungi. Preferences of lichenized fungi for very old (and mostly large) trees have been suggested to be due to continuity in substrate occurrence, which is important due to the slow growth and poor dispersal abilities of many species (Moe and Botnen 1997; Peck and McCune 1997). However, also the development of specic bark structures on old trees has been suggested to be a major reason for the preference for very old trees by several lichenized fungi (Gauslaa 1995). Many of these species have been assumed to be restricted to large trees situated in sunny sites in farmland landscapes. However, several of the species included in the present WKH study have also been found on relatively small, but very old oaks, on steep forested slopes (Ek et al. 1995). This suggests that these species might originally have been found also in natural sites and emphasises the importance of substrate continuity in relation to substrate quality. Of the studied factors, stand composition was classied as the least important (1% unique explainable variation) for occurrence of red-listed species. Thus, local habitat composition (see above) seemed to be much more important than stand composition for the occurrence of forest-associated red-listed bryophytes and fungi. In contrast,

1499 studies of birds and mammals in southern Sweden have suggested that adjacent habitats and stand composition on a relatively large scale are important for their occurrence (e.g. Hansson et al. 1995; Hansson 1997). Most WKHs are too small to harbour viable populations of many mammals and birds and management on a larger landscape scale is important for these organism groups. However, the WKH survey was not designed to investigate effects of surrounding landscapes on bryophytes and fungi (i.e. no large-scale landscape level variables were included). Thus, stand composition might be important also for these taxa, and furthermore, the long-term survival of red-listed species in WKHs might depend on the structure of surrounding areas. In conclusion, the present study quanties results from earlier studies suggesting that availability of high quality substrates (mainly old living trees and logs) is very important for the occurrence of red-listed bryophytes, lichenized and non-lichenized fungi (e.g. Berg et al. 1994; Rydin et al. 1997). Suitable substrates were generally found in old forests, but red-listed lichenized fungi were found in older forests than the other taxa. Sites with similar substrates, but situated in different regions (and climates), or with different ground moisture and nutrient conditions are needed to cover the full spectrum of habitat conditions suitable for different red-listed bryophytes and fungi. Most protected forest areas in Sweden consist of old forests, but the present forest nature reserve system covers 1.2% of the total forest area in southern Sweden, i.e. Gotaland and Svealand (Statistics Sweden 1999). Thus, many forest types (e.g. deciduous forests and spruce forests on nutrient-rich soils) are sparsely represented among nature reserves and national parks in Sweden (see also Nilsson and Gotmark 1992; Fridman 2000). Retention of trees and dead wood in the managed forest is of major importance for many red-listed species, since a sufcient increase in the protected area is unlikely. The relatively large amounts of dead wood in mature forests (Fridman and Walheim 2000) suggest that retention of snags and logs in silviculture is a possible measure in many areas, although active generation of dead wood might be needed in southern Sweden where dead wood decays faster (Fridman and Walheim 2000). Conservation measures in the relatively small WKHs, often surrounded by a matrix of managed forest, will therefore be of large importance. Nonetheless, monitoring of a selected number of WKHs, and investigations of reproduction and dispersal abilities of a spectrum of bryophyte and fungi species, would be desirable in order to evaluate the long-term quality of WKHs, not the least because the surroundings might affect survival and reproduction of red-listed species.

Acknowledgements Thanks to Lena Gustafsson, Nick Hodgetts and Per Milberg for comments on earlier versions of the paper. This study was funded by the Swedish University of Agricultural Sciences by the programme eld Analyses and Prediction of Biodiversity (to U.G.).

1500

Appendix
Number of occurrences at model sites, test sites and total number of occurrences for the 43 lichenized fungi species, nine bryophyte species and 11 non-lichenized fungi species included in the study. Abbreviated names are used in all gures. For more information on distribution, ecology and threats to 1998; Thor 1998. individual species see Larsson 1997; Hallingback Species Lichenized fungi ART SPAD BAC ROSE BIA MONA BIA SPHA BRY BICO BUE VIOL CAL LUCI CAL PARV CAL QUER CAT LAUR CHA CHLO CHA GRAC CHA PHAE CLA PARA CLI CORR COL SUBN EVE DIVA GYA FLOT GYA TRUN GYA ULMI LCA GLAB LCI EPIZ MEG GROS MEN TERE MIC AHLN NEP LAEV NOR PULC OPE ILLE OPE OCHR OPE SORE OPE VERM OPE VIRI PER MULT PHL AGEL PYR NITA RAM BALT SCH DECO SCH PERI SCL CONI SCL FARI SCL PERO SPH TURB a Arthonia spadicea Bacidiarosella Biatoridium monasteriense Biatora sphaeroides Bryoria bicolor Buellia violaceofusca Caloplaca lucifuga Calicum parvum Ca. quercinum Catinaria laureri Chaenotheca chlorella C. gracillima C. phaeocephala Cladonia parasitica Cliostomum corrugatum Collema subnigrescens Evernia divaricata G. otowii G. truncigena G. ulmi Lecanora glabrata B. chrysanta Megalaria grossa Menegazzia terebrata Microcalicium ahlneri Nephroma laevigatum Normandina pulchella Lecanographa amylacea Opegrapha ochrocheila O. sorediifera O. vermicellifera O. viridis Pertusaria multipuncta Phlyctis agelaea Pyrenula nitida Ramalina baltica Schismatomma decolorans Sc. pericleum Sclerophora coniophaea Sc. farinacea Sc. peronella Sphinctrina turbinata Model sites 123 95 21 94 17 77 107 20 42 33 96 35 427 20 307 78 32 52 47 283 229 25 207 71 35 53 72 103 25 31 31 48 44 114 470 71 191 212 77 24 34 39 Test sites 136 74 17 90 16 98 103 22 41 39 102 44 480 22 287 96 39 51 51 295 217 26 195 49 34 64 84 100 29 36 24 55 46 111 462 66 186 222 93 42 30 34 Total 259 169 38 184 33 175 210 42 83 72 198 79 907 42 594 174 71 103 98 578 446 51 402 120 69 117 156 203 54 67 55 103 90 225 932 137 377 434 170 66 64 73

1501 Appendix (continued )

Species USN FLOR Usnea orida Bryophytes ANAS HEL Anastrophyllum hellerianum BUXB VIR Buxbaumia viridis DICR DEN Dicranodontium denudatum DICR TAU Dicranum tauricum LOPH ASC Lophozia ascendens NECK PEN N. pennata NECK PUM Neckera pumila ORTH GYM Orthotrichum gymnostomum TRIC TOM Trichocolea tomentella Non-lichenized fungi ANT PULV Antrodia pulvinascens CLA PYXI Clavicorona pyxidata FIS HEPA Fistulina hepatica HOL MUCI Holwaya mucida INO HISP Inonotus hispidus PAC TUBE Pachykytospora tuberculosa PHE FERE Phellinus ferrugineus PHE FERF P. ferrugineofuscus PHE NIGL P. nigrolimitatus PHE POPU P. populicola SKE NIVE Skeletocutis nivea
a

Model sites 57 246 240 22 22 18 38 139 43 135 44 114 113 52 45 59 28 341 52 53 23

Test sites 57 254 245 18 27 15 31 144 52 140 53 102 81 41 55 64 15 343 60 49 27

Total 114 500 485 40 49 33 69 283 95 275 97 216 194 93 100 123 43 684 112 102 50

Not lichenized but grows on the lichenized fungus Pertusaria pertusa.

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