Forest Ecology and Management 315 (2014) 80–85

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Forest Ecology and Management

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Biodiversity of most dead wood-dependent organisms in thermophilic temperate oak woodlands thrives on diversity of open landscape structures ⇑ Jakub Horak a, , Stepan Vodka b, Jiri Kout c, Josef P. Halda d, Petr Bogusch e, Pavel Pech e a Czech University of Life Sciences Prague, Faculty of Forestry and Wood Sciences, Department of Forest Protection and Entomology, Kamycka 1176, CZ-165 21 Prague, Czech Republic b Czech Academy of Sciences, Institute of Entomology, Branisovska 31, CZ-370 05 Ceske Budejovice, Czech Republic c University of West Bohemia, Faculty of Education, Department of Biology, Geosciences and Environmental Education, Klatovska 51, CZ-306 19 Plzen, Czech Republic d Muzeum and Gallery of Orlicke Mts., Jiraskova 2, CZ-516 01 Rychnov nad Kneznou, Czech Republic e University of Hradec Kralove, Faculty of Science, Department of Biology, Rokitanskeho 62, CZ-500 03 Hradec Kralove, Czech Republic article info abstract

Article history: Oak and mixed deciduous forests with oaks are the most widespread woodland types in the central Received 17 July 2013 European lowlands. The aim of this study was to analyse how the biodiversity of saproxylic organisms Received in revised form 13 December 2013 (fungi, lichens, , and , bees and ) in thermophilic temperate oak woodlands respond Accepted 15 December 2013 to the openness in landscape structure of tree habitats. We sampled 32 sites in a split-plot design in Kri- Available online 11 January 2014 voklatsko (Czech Republic), which were chosen to include spatial diversity, including dense forests, open forests, woodland edges and solitary trees. A canonical correspondence analyses (CCA) and generalized Keywords: additive models (GAM) were used for analyses. The results indicated that the taxa studied showed differ- Lichens (Lichenes) ences in composition among the studied landscape structures and most taxa preferred more open Fungi Beetles (Coleoptera) and light conditions of the woodland environment. We also observed positive effect of the heterogeneity in open landscape structures on biodiversity of saproxylic organisms. As it is recently showed by ecolo- Saproxylic organisms gists, most of the thermophilic oak woodlands are threatened by succession, saproxylic organisms are Biodiversity management facing decline throughout the world and traditional forest management (e.g. game keeping, wood pastur- ing or coppicing) appears to be one solution to mitigate biodiversity loss. Ó 2013 Elsevier B.V. All rights reserved.

1. Introduction Woodlands dominated by oaks are recently facing the abandon- ment of traditional forest management, and the modern forest man- Many parts of the Earth are facing biodiversity loss and many agement of high forests (Peterken, 1993) together with artificial places important for their biological legacy are negatively affected regeneration are probably factors that have jeopardized their main- by the homogenisation and intensification of recent environmental tenance (Mielikainen and Hynynen, 2003). Wide areas of woodlands management methods (Vitousek et al., 1997; Dirzo and Raven, at lower altitudes have been pastured in the past by domestic ani- 2003). Many species are predicted to be lost before they have been mals (cattle, pigs, sheeps and goats), intensively coppiced or copp- explored for science, and in the best case scenario, specimens are at iced with standards, gathered for litter, used for game keeping and least currently preserved in several museum collections around the hunting and many disparate management types have created the world (Ponder et al., 2001; Horak et al., 2012). The number of fine mosaic of woodland patch types in which many species thrive threatened species is increasing and these species are losing their (Vera, 2000; Horak et al., 2012). The abandonment of such manage- former strongholds and survive on the edges of their distribution ment activities and the creation of conservation areas with unman- areas (Channell and Lomolino, 2000). Many formerly common spe- aged oaks have led to the expansion of early successional tree species cies and their habitats have been vanishing on various landscape such as birch, which shade lower canopy strata and overgrow soli- scales (Konvicka et al., 2008). However, in contrast, some species tary trees with limbs in the lower canopy (Ranius and Jansson, are increasing in abundance and distribution (e.g. extralimitally) 2000). A similar problem is observed with maples and ashes on high or are colonising new disjunctive areas (Kloot, 1984). Most proba- nutrient soils, beech at high altitudes on mesic (or deep) soils and ble causes are changes in land use and climate (Horak et al., with conifers (spruce and ) artificially planted under the oak 2013a). canopy (Hofmeister et al., 2004; Janik et al., 2011). The loss of formerly common species is reported also from tem- ⇑ Corresponding author. Tel.: +420 777838284. perate oak woodland areas (Buse et al., 2007; Hedl et al., 2010). E-mail address: [email protected] (J. Horak). Oak and mixed deciduous forests with oaks are the most

0378-1127/$ - see front matter Ó 2013 Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.foreco.2013.12.018 J. Horak et al. / Forest Ecology and Management 315 (2014) 80–85 81 widespread woodland habitats in the central European lowland Only species with an affinity for dead wood habitats (hereafter and adjacent areas (Svoboda et al., 2011). In the temperate climatic called saproxylics) were evaluated. For species, all obligate zone, various types of oak woodlands naturally occur up to ca. saproxylics were taken into account (Schmidl and Bußler, 2004). 600 m a.s.l., especially on dry and semi-dry sites (Neuhauslova, All dead wood-dependent fungal species (Hagara et al., 2005) were 1998). The majority of European oak species are sun-loving trees, analysed, except those with corticioid fruiting bodies; pyrenomy- whose recruitment is dependent on favourable micro-climatic con- cetes and inoperculate discomycetes with small apothecia were ditions during the fruiting stage of mature trees and for seedling also not sampled. Most saproxylic species of hymenopterans use growth and development (Rolecek, 2005; Ugurlu et al., 2012). Oaks dead wood for nesting and some are able to build nests in other are also known for their stamina and high quality wood and are substrates, thus, we only evaluated species with a predominant often protected as single specimens or monument trees. dead wood strategy (Macek et al., 2010). The associations of lichens Furthermore, acorns are nutrient-rich feed for domestic and semi- with dead wood substrates in central Europe are more complicated domestic such as pigs or wild boars (Vera, 2000). Veteran than for the other studied taxa (Jansova and Soldan, 2006). As oaks are one of the most suitable hosts for organisms that are depen- lichens are mostly long-lived organisms, it is often hard to deter- dent on dead wood (Svoboda et al., 2011; Alexander, 2013). mine whether a species is associated only with a living tree (i.e. Organisms dependent on dead wood, like beetles and fungi, arboricolous) or is more opportunistic and dependent only on have recently been the focus of forest ecology (Lindhe et al., substrates generated by wood plants. Therefore, we used all lichen 2005; Buse et al., 2007; Norden et al., 2008). These organisms are species in our analyses that were found on dead and partly living highly interconnected with woodland habitat types and many are wood, because most trees in old-growth forests include many dead bioindicators (Svoboda et al., 2010). Studies performed at the spe- wood microhabitats (Winter and Moller, 2008). cies level have shown that the openness of the surrounding wood- land environment often leads to an increase in species population 2.3. Sampling of taxa density (Buse et al., 2007). A similar situation is observed in studies at guild or family levels, where most taxa thrive on canopy open- Fungi and lichens were sampled using circular plots with a ness (Lindhe et al., 2005; Horak and Rebl, 2013). Although a gap re- radius of 20 m surrounding the traps (area 1256 m2). For solitary mains in the knowledge of multi-taxon responses, and non-boreal trees, the shape of the sampling plot was adapted to the tree- landscapes are under-represented with respect to the number of dominated habitat in the surroundings, but also had a fixed total studies. Namely, species diversity is much higher in temperate area. The centre of the sampling plot was 20 m from the edge of and Mediterranean woodlands, and oak-dominated or mixed the woodland for the edge category. Fungi were sampled four woodlands are European hot-spots of biodiversity (Blasi et al., times in June, September and October 2011 and May 2012. Lichens 2010; Buse et al., 2010). Solitary oak trees are known to be very were sampled once during August 2011. Beetles and hymenopter- rich with respect to the number of species of disparate organisms ans were sampled using large window trunk tree traps (Horak, (Svoboda et al., 2010). Nevertheless, the responses of dead wood- 2011), which are known to be highly effective for trapping dead dependent organisms to environmental changes can be of high wood associates including flightless fauna (Horak et al., 2013b). interest for suitable biodiversity management activities. The traps were active and regularly emptied during the 2011 veg- etation season (May–September). The aim of this study was to analyse how the biodiversity of saproxylic organisms responds to the openness and diversity in 2.4. Study environmental variables and their sampling landscape structure of tree habitats in thermophilic temperate oak woodlands, using a multi-taxa approach. Our study focused on four main landscape structures of tree Namely, we focused on (i) species composition, (ii) richness and habitats: (iii) its trends of four taxa (fungi, lichens, beetles, and ants, bees and wasps) and their influencing by four landscape structures (i) solitary trees (canopy openness: mean = 32.51% ± 4.57 SE) (solitary trees, woodland edges, open forests and dense forests). with mean distance of 94.75 m (SE = 22.22 m) from the near- est forest edge and high coverage of plants and shrubs in the understorey, 2. Methods (ii) natural woodland edges (32.05% ± 5.92 SE) between forest and non-forest area (grasslands and arable land) with med- 2.1. Study area ium coverage of plants and shrubs in the understorey, (iii) forests with open canopy conditions (32.87% ± 4.05 SE) and Krivoklatsko is situated in the western part of the Central medium coverage of plants and low coverage of shrubs in Bohemian range about 20 km west of Prague (coordinates: the understorey, 50.00N; 13.88E). It is known within the Czech Republic for a high (iv) dense forests with closed canopy conditions (12.88% ± 0.68 biodiversity, which has recently been highlighted by attempts to SE) with low coverage of plants and absence of shrubs in protect this area as a national park (Hula, 2009). Its large area the understorey. (ca. 630 km2) of low and medium altitudes (ca. 400 m a.s.l.) contains diversified land use, geomorphology and a large number First three landscape structures (i–iii) reflected different open of disparate nature conservation and cultural heritage areas, such canopy conditions and one (iv) reflected closed canopy conditions. as Krivoklatsko UNESCO Biosphere Reserve and Krivoklatsko In total, we sampled 32 sites that were designed as split plots – Protected Landscape Area, with nearly 30 small-area protected i.e. eight whole plots contained all four landscape structures. Three sites. Krivoklatsko is one of the most forested parts of the Czech pairs of whole plots were distributed in or near three protected Republic, with 62% covering by forests (Hula, 2009). areas recently listed (www.pralesy.cz) as the most natural wood- lands in the Czech Republic (Brdatka, Nezabudicke skaly and Na 2.2. Study taxa Babe) and two pairs between protected areas Jouglovka and Kohoutov. The main reason for this choice was natural or semi- We studied four taxa: fungi (Fungi), lichens (Lichenes), beetles natural tree species composition of above mentioned protected (Coleoptera) and ants, bees and wasps (Hymenoptera: Aculeata). areas and their surroundings (Nemec and Lozek, 1996). 82 J. Horak et al. / Forest Ecology and Management 315 (2014) 80–85

Coordinates (latitude and longitude) of the plot centres as the spatial terms were measured directly in the field using a Garmin Colorado 300. The amount of dead wood as the best reflections of habitat area in woodland habitats for saproxylic organisms was measured in each plot. All dead wood (mean = 1.20; SE = 0.26; min = 0.01; max = 4.60 m3) was measured, including all lying and standing pieces to 5 m above the ground and with a diameter >0.10 m with- in a 20 m radius surrounding the traps. When only thinner diame- ter dead wood was found, we used a value of 0.01 m3. Canopy openness (mean = 27.58; SE = 2.54; min = 10.36; max = 61.09%) was measured using a Nikon COOLPIX 995 camera with a Nikon FC-E8 Fish Eye converter.

2.5. Data analyses Fig. 1. Comparison of results of canonical correspondence analyses (CCA) on species composition of the studied taxa (fungi, lichen, Coleoptera, and Hymenop- tera) in the thermophilic oak woodland area. Independent contribution of the All analyses were performed in CANOCO. We used a presence/ studied categories (solitary trees, woodland edges, open forests and dense forests) absence species matrix as it gave a better final comparison of taxa is white and joint contribution with space and dead wood as co-variables is black. sampled using different techniques (Horak et al., 2013c). A canon- F and P values of CCA are indicated above joint contributions. ical correspondence analysis (CCA) was used as the most appropri- ate method of multivariate statistics. Landscape structures were solitary trees (Fig. 2c), whereas hymenopterans showed the high- used as categorical variables and dead wood with coordinates est preference for ecotonal habitats and the species composition and their crossed and quadratic products were treated as continu- here was most distinct from that of other habitats (Fig. 2d). ous co-variables (Horak, 2013). A global permutation test was set The species richness isolines did not illustrate any clear trend at 9,999 permutations under the full model. The permutation type with respect to the studied design for fungi, although the samples was restricted for a split-plot design with the whole plot freely from the edges and dense forest habitats showed a bit higher spe- exchangeable. Due to the presence of empty species samples for cies richness than other habitats (Fig. 3a), which is in agreement fungi, lichens and Hymenoptera, we used the virtual opportunistic with previous results (Fig. 2a). The species richness of lichens species present in all samples. The potential bias of multi-colinearity and beetles decreased from solitary trees towards closed forests was detected using the variance inflation factor (VIF). The joint ef- (Fig. 3b and c). Open forests and edges possessed the highest fect of the studied categories of landscape structures with co-vari- species richness of saproxylic Hymenoptera (Fig. 3d). ables was computed using the process of variance partitioning. The Trace/Total interia ratio was used for comparison of the CCA 4. Discussion among the studied taxa. Generalized additive models (GAM) were used to compute and visualise species richness gradients for each The results showed that most taxa preferred diversification of studied taxa in CanoDraw. The Akaike information criterion (AIC) the more open and light conditions of the woodland environment. was used to select the best response of the species richness (num- Highly diversified open canopy structures like solitary trees, natu- ber of species as a dependent variable) to the studied categories. ral edges and open forests are potentially primary objectives of the All photographs of canopy were evaluated using a Gap Light present forest management (Salek et al., 2013). Furthermore, some Analyser 2.0. The difference in canopy openness between dense saproxylics, namely beetles and lichens, preferred solitary trees, and open forests was assessed by paired t test. which are usually not considered as a forest habitat. This might lead to the conclusion that some saproxylic organisms do not pre- 3. Results fer forest. Although, the response of the studied taxa might be influenced by the fact that open thermophilic oak woodlands in We observed 78 species of saproxylic fungi, 36 species of our study area have a high temporal continuity (Neuhauslova, lichens, 153 species of beetles and 32 species of hymenopterans 1998). Oak woodlands have a naturally higher canopy openness (Tables S1–S4). (Rolecek, 2005; Ugurlu et al., 2012), due to the requirements of The control analysis showed that there was a significant differ- the dominating tree species (Quercus spp.) – associated organisms ence in canopy openness between dense and open forests are thus more adapted to light conditions in woodlands. (t = À5.21; P < 0.01). Except for the natural fauna and flora, open canopies promote The effect of the studied landscape structures (solitary trees, the presence of species adapted to warm climatic conditions woodland edges, open forests and dense forests) on species (Ugurlu et al., 2012; Alexander, 2013). This is well-illustrated by composition of all studied taxa was significant and independent the occasional occurrence of the Mediterranean long-horned bee- contribution was nearly the same for all taxa. Lichens showed tle, Purpuricenus kaehleri (L.) in the Czech Republic. This species the highest independent and joint contributions and beetles shared is not threatened in Europe (Nieto and Alexander, 2010) and in the lowest contribution joint with co-variables (Fig. 1). the Czech Republic, is mainly distributed in the two most thermo- Most of the studied taxa showed a distinct species composition philic areas: Krivoklatsko and southern Moravia (Slama, 1998). The among the studied landscape structures (e.g. fungi in dense forests species was considered in the Red-list as regionally extinct (Farkac and edges; Fig. 2a). The results also showed that the species et al., 2005), but was once again collected in our study area in pres- composition of most taxa was almost disparate in dense forests, ent years, probably due to favourable climate conditions. compared to other habitats (Fig. 2a–c), and were also poorer in Oak woodlands have often been managed in forest systems number of species in most cases (Fig. 2). Fungal communities were known for higher canopy openness, such as coppices, coppices most similar in open forests and solitary trees (Fig. 2a). The species with standards or pasture woodlands (Vera, 2000; Horak et al., richness of lichens appeared to increase with an increasing gradi- 2012; Fartmann et al., 2013). Currently, there are only a few ent of openness in landscape structures (Fig. 2b). Beetles preferred over-matured stands in the Czech Republic, which are indicative J. Horak et al. / Forest Ecology and Management 315 (2014) 80–85 83

Fig. 2. Sample-environmental attribute plots (a high circle radius shows a high species richness of samples) for the studied taxa: (a) fungi, (b) lichens, (c) Coleoptera and (d) Hymenoptera in the thermophilic temperate oak woodland area. Sub-graphs in the upper right corners show the value of the weighted species mean in each studied category. First and second CCA axes are shown. of former coppice management and the only recently active cop- only recent effect of conservation management is to leave the for- pice-like stands are riparian corridors dominated by alders (Pecha, ests unmanaged (Hort et al., 1999) – however, this conservation 2010). Except for presence of coppicing and pasturing, one of the activity mostly leads to canopy closure (Horak et al., 2012) and important factors for high biodiversity might be the absence of to the dominance of late successional tree species such as beech, modern forest management (with the artificial regeneration of especially in thermophilic oak forests on deep soils. Non-interven- stands after harvesting), which causes a dominance of oak in veg- tion management in thermophilic oak woodlands mostly leaves etative and natural regeneration and also leads to a sparser canopy open structures only in stands on slopes, with shallow rocky soils, caused by damage by high game stocks on natural regeneration. In a lower nutrition quality and a humus layer caused by natural our study area, historical sources also mention a high diversity of processes such as erosion (Rolecek, 2005) and by a higher accumu- other, so-called traditional forest management activities – such lation of game due to less activity from hunters. as litter gathering or basketry, which also promote an open forest structure (Horak et al., 2012). 4.1. Management possibilities The high area of the Krivoklatsko woodlands was managed in the past for game hunting, which has been carried out since medi- Thermophilic oak forests on deep soils represent successionally eval times – i.e. a vast area was fenced in as a game park for the unstable vegetation created by man (Svoboda, 1943; Rolecek, Czech king and nobility (Horak and Rebl, 2013). The recent hunting 2005). Thus, their survival and spatial structure is dependent on community has been criticized for high game stocks, which cause active forest management. Traditional management activities, such damage to natural regeneration and timber production (Kupcak, as coppicing or wood pasturing, represent one solution (Hedl et al., 2005). However, it is known that a high game stock pressure is a 2010; Hartel et al., 2013) – however, these have been highly prob- successful biological agent that acts against forest succession and lematic in recent times, as mentioned above. towards closed canopy woodlands dominated by beech and coni- Game parks, such as Lany Game Park within the studied area, fers (Janik et al., 2011). with its large area of 30 km2, represent one management possibil- The main problem of closing canopies and following biodiver- ity towards open canopy woodlands (Horak and Rebl, 2013). There sity loss, is that nearly all traditional forest management types are also many ways to manage commercial stands with high forest, are restricted (e.g. wood pasturing) or are affected by Czech law which could (at least ephemerally) mimic the openness of forest (e.g. coppicing; Utinek, 2004). Except for commercial forestry, the management – these are larger areas with prolonged shelter-wood

Table 1 Species richness trends in the thermophilic oak woodland. Results of GAM using selection based on AIC, significant P values are in bold.

Taxa First CCA axis DF Second CCA axis DF Resid. DF FP AIC Fungi 1 6 24 4.54 <0.01 299.16 Lichen 1 4 26 2.61 <0.05 263.21 Coleoptera 4 4 23 2.78 <0.05 2472.58 Hymenoptera 1 6 24 3.44 <0.05 93.63 84 J. Horak et al. / Forest Ecology and Management 315 (2014) 80–85

Fig. 3. Response of particular taxa: (a) fungi, (b) lichens, (c) Coleoptera and (d) Hymenoptera to the studied categories (black) in the thermophilic temperate oak woodland area. Grey isolines reflect the trend (not total value) in species richness derived from GAM (Table 1); the first and second CCA axes are shown. cuttings, leaving several trees (more damaged = more effective) in vegetation created by man (Lozek, 1973), without active forest clear-cuts, or the maintenance, promotion and planting of solitary management, they often succumb to the expansion of late succes- individuals or avenues in grasslands, arable land and along roads. sional trees (Rolecek, 2005). The biodiversity of the studied taxa, in The worst solution for the protection of regional biodiversity of most cases, is driven by openness and diversity in the landscape thermophilic oak woodlands appears to be to leave the stands structures of thermophilic oak woodlands. There are many rare unmanaged. This solution, if successfully practiced in beech wood- or threatened saproxylic organisms connected with oak tree habi- lands (Gossner et al., 2013), is applicable only to dry forests on tats (Buse et al., 2007; Norden et al., 2008; Svoboda et al., 2011; shallow rocky soils, which are naturally disturbed by soil erosion, Petrakova and Schlaghamersky, 2011). Saproxylic organisms are or semi-naturally by game stocks. in decline throughout the Europe (Nieto and Alexander, 2010) and thermophilic oak woodlands, as one of their main habitats, are threatened by succession (Moga et al., 2009). Thus, active forest 5. Conclusions management that causes openness and diversification in the management of oak woodlands is one of the solutions to mitigate The first topic that needs to be addressed is the restoration of biodiversity loss. traditional forest management systems such as coppicing and wood-pasturing, at least in conservation areas. This is because Acknowledgements these management types retain the open structures and avoid the dominance of late successional tree species, namely beech, We would like to thank P. Stloukal for support, K. Rébl for the which has lower commercial potential than oak and furthermore, identification of beetles, D. Romportl for fish eye equipment, the low commercial potential of coppice-like management appears Mr. Besitzer for correction of English and two anonymous referees to be a pseudo-issue in recent times (Utinek, 2004; Kupcak, 2010). provided constructive comments. The second goal is the maintenance of high game stocks, even when they are distributed outside the game parks. Whereas wood Appendix A. Supplementary material pasturing is prohibited within the Czech Republic, the only way to mimic this formerly widely used management method is via the Supplementary data associated with this article can be found, maintenance of high game stocks, especially in conservation areas, in the online version, at http://dx.doi.org/10.1016/j.foreco.2013. where their damage has a low commercial effect. Furthermore, 12.018. game hunting is known to increase local incomes and improve human diet (Hawkes et al., 1991; Wilkie and Carpenter, 1999). The third aim is support for local activities towards the mainte- References nance and restoration of solitary trees in non-woodland land- Alexander, K., 2013. Ancient trees, grazing landscapes and the conservation of scapes, an activity that is often performed by non-governmental deadwood and wood decay invertebrates. In: Rotherham, I.D. (Ed.), Trees, organisations. Forested Landscapes and Grazing Animals, pp. 330–337. Semi-natural oak woodlands are one of the most widespread Blasi, C., Marchetti, M., Chiavetta, U., Aleffi, M., Audisio, P., Azzella, M.M., Burrascano, S., 2010. Multi-taxon and forest structure sampling for forest types in the lowlands of Central Europe (Svoboda et al., identification of indicators and monitoring of old-growth forest. Plant Biosyst. 2011), although since they represent successionally unstable 144, 160–170. J. Horak et al. / Forest Ecology and Management 315 (2014) 80–85 85

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Effect of site level environmental variables, spatial autocorrelation and tree size on saproxylic beetles associated with old oaks. Biol. Conserv. 95, and sampling intensity on communities in an ancient temperate 85–94. lowland woodland area. PLoS One 8, e81541. Rolecek, J., 2005. Vegetation types of dry-mesic oak forests in Slovakia. Preslia 77, Horak, J., Rebl, K., 2013. The species richness of click beetles in ancient pasture 241–261. woodland benefits from a high level of sun exposure. J. Insect Conserv. 17, 307– Salek, L., Zahradnik, D., Marusak, R., Jerabkova, L., Merganic, J., 2013. Forest edges in 318. managed riparian forests in the eastern part of the Czech Republic. For. Ecol. Horak, J., Chobot, K., Horakova, J., 2012. Hanging on by the tips of the tarsi: a review Manage. 305, 1–10. of the plight of the critically endangered saproxylic beetle in European forests. J. Schmidl, J., Bußler, H., 2004. Ökologische Gilden xylobionter Käfer Deutschlands. Nat. Conserv. 20, 101–108. 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Sci. 50, 38–46. and hornbeam populations along a 4-m elevation gradient in an alluvial forest. Vera, F.W.M., 2000. Grazing Ecology and Forest History. CABI. Eur. J. For. Res. 130, 197–208. Vitousek, P.M., Mooney, H.A., Lubchenco, J., Melillo, J.M., 1997. Human domination Jansova, I., Soldan, Z., 2006. The habitat factors that affect the composition of of Earth’s ecosystems. Science 277, 494–499. bryophyte and lichen communities on fallen logs. Preslia 78, 67–86. Wilkie, D.S., Carpenter, J., 1999. The potential role of safari hunting as a source of Kloot, P.M., 1984. The introduced elements of the flora of southern Australia. J. revenue for protected areas in the Congo Basin. Oryx 33, 340–345. Biogeogr. 11, 63–78. Winter, S., Moller, G.C., 2008. Microhabitats in lowland beech forest as monitoring Konvicka, M., Benes, J., Cizek, O., Kopecek, F., Konvicka, O., Vitaz, L., 2008. How too tool for nature conservation. For. Ecol. Manage. 255, 1251–1261. much care kills species: grassland reserves, agrienvironmental schemes and Supporting information

Biodiversity of most dead wood‐dependent organisms in thermophilic temperate oak woodlands thrives on diversity of open landscape structures

Jakub Horaka,*, Stepan Vodkab, Jiri Koutc, Josef P. Haldad, Petr Bogusche, Pavel Peche

aCzech University of Life Sciences Prague, Faculty of Forestry and Wood Sciences, Department of Forest Protection and Entomology, Kamycka 1176, CZ‐165 21 Prague, Czech Republic bCzech Academy of Sciences, Institute of Entomology, Branisovska 31, CZ‐370 05 Ceske Budejovice, Czech Republic cUniversity of West Bohemia, Faculty of Education, Department of Biology, Geosciences and Environmental Education, Klatovska 51, CZ‐306 19 Plzeň, Czech Republic dMuzeum and Gallery of Orlicke Mts., Jiraskova 2, CZ‐516 01 Rychnov nad Kneznou, Czech Republic eUniversity of Hradec Kralove, Faculty of Science, Department of Biology, Rokitanskeho 62, CZ‐500 03 Hradec Kralove, Czech Republic

*Corresponding author: E‐mail address: [email protected] (J. Horak).

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Table S1

A check‐list of fungi.

Species Number of plots Aleurodiscus disciformis (DC) Pat. 1 Amylostereum areolatum (Chaillet ex Fr.) Boidin 1 Antrodiella semisupina (Berk. & M.A. Curtis) Ryvarden 1 Auricularia auricula‐judae (Bull.) Quél. 6 Basidioradulum radula (Fr.) Nobles 2 Biscogniauxia nummularia (Bull.) Kuntze 2 Bjerkandera adusta (Willd.) P. Karst. 1 Byssomerulius corium (Pers.) Parmasto 3 Calocera cornea (Batsch) Fr. 1 Cinereomyces lindbladii (Berk.) Jülich 5 Clitopilus hobsonii (Berk. & Broome) P.D. Orton 2 Colpoma sp.1 1 Coriolopsis gallica (Fr.) Ryvarden 1 Crepidotus sp.1 1 Dacrymyces stillatus Nees 3 Daedalea quercina (L.) Pers. 7 Daedaleopsis confragosa (Bolton) J. Schröt. 1 Datronia mollis (Sommerf.) Donk 2 Diplomitoporus flavescens (Bres.) Domański 1 Exidia cartilaginea Lund. & Neuh. 1 Exidia nigricans (With.) P. Roberts 9 Exidia truncata Fr. 2 Fomitopsis pinicola (Sw.) P. Karst. 1 Ganoderma applanatum (Pers.) Pat. 1 Hapalopilus nidulans (Fr.) P. Karst. 2 Hericium erinaceus (Bull.) Pers. 1 Hymenochaete rubiginosa (Dicks.) Lév. 3 Hypholoma fasciculare (Huds.) P. Kumm. 1 Hypoxylon howeanum Peck 4 Inonotus dryophilus (Berk.) Murrill 1 Lentinellus ursinus (Fr.) Kühner 1 Lenzites gibbosa (Pers.) Hemmi 1 Lopharia spadicea (Pers.) Boidin 1 Marasmius rotula (Soop.) Fr. 3 Mycena crocata (Schrad.) P. Kumm. 1 Mycena galericulata (Scop.) Gray 6 Mycena polygramma (Bull.) Gray 2

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Species Number of plots Mycetinis scorodonius (Fr.) A.W. Wilson & Desjardin 2 Nectria cinnabarina (Tode) Fr. 2 Panellus stipticus (Bull.) P. Karst. 1 Phaeolus schweinitzii (Fr.) Pat. 2 Phellinus contiguus (Pers.) Pat. 15 Phellinus ferruginosus (Schrad.) Pat. 1 Phellinus robustus (P. Karst.) Bourdot & Galzin 4 Phellinus tuberculosus (Baumg.) Niemelä 1 Pleurotus pulmonarius (Fr.) Quél. 1 Plicaturopsis crispa (Pers.) D.A. Reid 1 Pluteus cervinus (Schaeff.) P. Kumm. 2 Pluteus leoninus (Schaeff.) P. Kumm. 1 Pluteus plautus (Weinm.) Gillet 1 Pluteus pouzarianus Singer 1 Polyporus arcularius (Batsch) Fr. 1 Polyporus ciliatus Fr. 1 Postia caesia (Schrad.) P. Karst. 1 Postia guttulata (Peck ex Sacc.) Jülich 2 Radulomyces molaris (Chaillet ex Fr.) M.P. Christ. 13 Schizophyllum commune Fr. 3 Schizopora flavipora (Berk. & M.A. Curtis ex Cooke) Ryvarden 9 Schizopora radula (Pers.) Hallenb. 14 Serpula himantioides (Fr.) P. Karst. 3 Skeletocutis nivea (Jungh.) Jean Keller 1 Steccherinum bourdotii Saliba & A. David 2 Steccherinum fimbriatum (Pers.) J. Erikss. 2 Steccherinum ochraceum (Pers.) Grey 3 Stereum gausapatum (Fr.) Fr. 3 Stereum hirsutum (Willd.) Pers. 20 Stereum rameale (Pers.) Burt 3 Stereum rugosum Pers. 4 Trametes hirsuta (Wulfen) Lloyd 1 Trametes versicolor (L.) Lloyd 3 Tremella encephala Willd. 1 Tremella mesenterica Retz. 1 Trichaptum abietinum (Dickson) Ryvarden 2 Vuilleminia sp.1 3 Xerula radicata (Relhan) Dörfelt 1 Xylaria hypoxylon (L.) Grev 6 Xylaria longipes Nitschke 1 Xylaria polymorpha (Pers.) Grev. 1

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Table S2

A check‐list of lichens.

Species Number of plots Amandinea punctata (Hoffm.) Coppins et Scheid. 2 Anisomeridium polypori (Ellis et Everh.) M. E. Barr 1 Bacidina sulphurella (Samp.) M. Hauck et V. Wirth 3 Chaenotheca ferruginea (Turner et Borrer) Mig. 4 Cladonia coniocraea (Flörke) Spreng. 2 Cladonia digitata (L.) Hoffm. 2 Cladonia fimbriata (L.) Fr. 1 Cladonia macilenta Hoffm. 1 Coenogonium pineti (Schrad. ex Ach.) Lücking et Lumbsch 8 Evernia prunastri (L.) Ach. 1 Flavoparmelia caperata (L.) Hale 1 Hypocenomyce scalaris (Ach.) M. Choisy 12 Hypogymnia physodes (L.) Nyl. 8 Imshaugia aleurites (Ach.) S. L. F. Mey. 1 Lecanora conizaeoides Nyl. ex Cromb. 14 Lecanora expallens Ach. 8 Lepraria incana (L.) Ach. 21 Melanelixia fuliginosa (Fr. ex Duby) O. Blanco et al. 4 Micarea micrococca (Körb.) Gams ex Coppins 7 Parmelia sulcata Taylor 8 Parmeliopsis ambigua (Wulfen) Nyl. 2 Pertusaria leioplaca DC. 1 Phaeophyscia orbicularis (Neck.) Moberg 1 Physcia adscendens (Fr.) H. Olivier 1 Physcia dubia (Hoffm.) Lettau 1 Physcia tenella (Scop.) DC. 5 Placynthiella icmalea (Ach.) Coppins et P. James 1 Platismatia glauca (L.) W. L. Culb. et C. F. Culb. 1 Porina aenea (Wallr.) Zahlbr. nom. illeg. 2 Pseudevernia furfuracea (L.) Zopf 1 Punctelia subrudecta (Nyl.) Krog 1 Scoliciosporum chlorococcum (Graewe ex Stenh.) Vězda 5 Trapeliopsis flexuosa (Fr.) Coppins et P. James 1 Trapeliopsis pseudogranulosa Coppins et P. James 1 Xanthoria candelaria (L.) Th. Fr. 2 Xanthoria parietina (L.) Th. Fr. 2

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Table S3

A check‐list of beetles.

Species Number of plots Abraeus granulum Erichson, 1839 1 Allecula morio (Fabricius, 1787) 3 Allecula rhenana Bach, 1856 1 Alosterna tabacicolor (DeGeer, 1775) 1 nigroflavus (Goeze, 1777) 2 Ampedus pomorum (Herbst, 1784) 4 Ampedus rufipennis (Stephens, 1830) 2 Ampedus sinuatus Germar, 1844 2 Anaesthetis testacea (Fabricius, 1781) 2 Anaglyptus mysticus (Linnaeus, 1758) 3 Anaspis flava (Linnaeus, 1758) 4 Anisotoma humeralis (Fabricius, 1792) 1 Aplocnemus impressus (Marsham, 1802) 1 Bitoma crenata (Fabricius, 1775) 1 Brachygonus megerlei (Lacordaire, 1835) 6 Cardiophorus gramineus (Scopoli, 1763) 1 Cerylon ferrugineum Stephens, 1830 11 Cerylon histeroides (Fabricius, 1792) 1 Cis boleti (Scopoli, 1763) 2 Cis micans (Fabricius, 1792) 1 Cis pygmaeus (Marsham, 1802) 1 Clytus tropicus (Panzer, 1795) 1 Conopalpus testaceus (Olivier, 1790) 3 Corticaria alleni Johnson, 1974 1 Corticaria bella L. Redtenbacher, 1849 1 Corticaria longicornis (Herbst, 1793) 1 Corticeus linearis (Fabricius, 1790) 1 Corticeus unicolor Piller & Mitterpacher, 1783 1 Cortodera humeralis (Schaller, 1783) 2 Crepidophorus mutilatus (Rosenhauer, 1847) 1 Cryptophagus fuscicornis Sturm, 1845 1 Cryptophagus labilis Erichson, 1846 2 Cryptophagus micaceus Rey, 1889 6 Cyrtanaspis phalerata (Germar, 1831) 3 Dacne bipustulata (Thunberg, 1781) 16 Dasytes aeratus Stephens, 1829 4 Dasytes niger (Linnaeus, 1761) 1

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Species Number of plots Dasytes plumbeus (O.F. Müller, 1776) 7 Dasytes virens (Marsham, 1802) 2 punctatus (Herbst, 1792) 1 linearis (Linnaeus, 1758) 2 chrysomelina Sturm, 1837 5 Dorcatoma flavicornis (Fabricius, 1792) 1 villosus (Fabricius, 1792) 5 Endomychus coccineus (Linnaeus, 1758) 1 Enedreytes sepicola (Fabricius, 1792) 2 Enicmus atriceps Hansen, 1962 3 Ennearthron cornutum (Gyllenhal, 1827) 3 guttata (Olivier, 1790) 1 Ernoporicus fagi (Fabricius, 1798) 4 Ernoporus tiliae (Panzer, 1793) 1 Eucnemis capucina Ahrens, 1812 7 Euglenes pygmaeus (De Geer, 1774) 1 Gastrallus immarginatus (P. W. J. Müller, 1821) 3 Glischrochilus quadrisignatus (Say, 1835) 1 Gnorimus nobilis (Linnaeus, 1758) 1 Grammoptera ruficornis (Fabricius, 1781) 1 Hadrobregmus pertinax (Linnaeus, 1758) 1 costatus (Gene, 1830) 1 Hesperus rufipennis (Gravenhorst, 1802) 6 Hylis foveicollis (C.G. Thomson, 1874) 1 Ischnomera cyanea (Fabricius, 1787) 2 Isoriphis marmottani (Bonvouloir, 1871) 2 Leiopus nebulosus (Linnaeus, 1758) 4 Lissodema denticolle (Gyllenhal, 1813) 1 Litargus connexus (Fourcroy, 1785) 9 Lucanus cervus (Linnaeus, 1758) 2 Magdalis flavicornis (Gyllenhal, 1836) 2 Malachius bipustulatus (Linnaeus, 1758) 1 Megatoma undata (Linnaeus, 1758) 3 castanipes (Paykull, 1800) 2 Melanotus crassicollis (Erichson, 1841) 7 Melanotus villosus (Fourcroy, 1785) 24 Melasis buprestoides (Linnaeus, 1761) 1 Molorchus minor (Linnaeus, 1758) 1 humeralis (Linnaeus, 1758) 2 Mordellochroa abdominalis (Fabricius, 1775) 1 Mycetochara axillaris (Paykull, 1799) 3 Mycetochara maura (Fabricius, 1792) 10

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Species Number of plots Mycetophagus piceus (Fabricius, 1787) 3 Mycetophagus populi Fabricius, 1798 2 Mycetophagus quadriguttatus P.W. J. Müller, 1821 2 Mycetophagus quadripustulatus (Linnaeus, 1761) 2 Nemozoma elongatum (Linnaeus, 1761) 2 Oberea linearis (Linnaeus, 1761) 1 Oligomerus brunneus (Olivier, 1790) 7 mollis (Linnaeus, 1758) 10 cinerea (Mulsant, 1839) 2 Orchesia fasciata (Illiger, 1798) 2 Orchesia micans (Panzer, 1795) 3 Orchesia minor Walker, 1837 2 Orchesia undulata Kraatz, 1853 1 Osphya bipunctata (Fabricius, 1775) 1 Oxylaemus cylindricus (Creutzer in Panzer, 1796) 3 Paromalus flavicornis (Herbst, 1792) 5 testaceus (Hellwig, 1792) 2 Phloiotrya rufipes (Gyllenhal, 1810) 5 Phymatodes rufipes (Fabricius, 1776) 2 Phymatodes testaceus (Linnaeus, 1758) 8 Pityogenes chalcographus (Linnaeus, 1761) 1 Plagionotus detritus (Linnaeus, 1758) 2 Platydema violacea (Fabricius, 1790) 1 Platypus cylindrus (Fabricius, 1792) 3 albinus (Linnaeus, 1758) 2 Prionocyphon serricornis (P.W. & J. Müller, 1821) 3 Prionychus ater (Fabricius, 1775) 1 Procraerus tibialis (Lacordaire in Boisduval & Lacordaire, 1835) 2 Protaetia marmorata (Fabricius, 1792) 1 Pseudocistela ceramboides (Linnaeus, 1758) 1 Ptilinus pectinicornis (Linnaeus, 1758) 5 rufipes Olivier, 1790 1 Quedius xanthopus Erichson, 1839 8 Rhagium mordax (DeGeer, 1775) 3 Rhamnusium bicolor (Schrank, 1781) 3 Rhizophagus bipustulatus (Fabricius, 1792) 7 Rhyncolus ater (Linnaeus, 1758) 1 Ropalopus femoratus (Linnaeus, 1758) 2 Salpingus planirostris (Fabricius, 1787) 3 Salpingus ruficollis (Linnaeus, 1761) 5 Scaphidium quadrimaculatum Olivier, 1790 1 Scolytus carpini (Ratzeburg, 1837) 8

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Species Number of plots Scolytus intricatus (Ratzeburg, 1837) 10 Scolytus mali (Bechstein, 1805) 1 Scolytus rugulosus (P.W. J. Müller, 1818) 1 Sepedophilus testaceus (Fabricius, 1792) 1 Silvanus unidentatus (Fabricius, 1792) 2 Stenagostus rhombeus (Olivier, 1790) 3 Stenichnus godarti (Latreille, 1806) 2 Stenomax aeneus (Scopoli, 1763) 5 dubia (Laicharting, 1784) 1 Stereocorynes truncorum (Germar, 1824) 3 Synchita humeralis (Fabricius, 1792) 3 Synchita separanda (Reitter, 1881) 1 starkii Chevrolat, 1859 4 Thanasimus formicarius (Linnaeus, 1758) 2 Tillus elongatus (Linnaeus, 1758) 2 bucephala (Costa, 1854) 4 Trichoceble memnonia (Kiesenwetter, 1861) 2 Triplax aenea (Schaller, 1783) 1 Triplax lepida Faldermann, 1835 1 Triplax russica (Linnaeus, 1758) 3 Trypodendron lineatum (Olivier, 1795) 1 Uleiota planata (Linnaeus, 1761) 1 Velleius dilatatus (Fabricius, 1787) 2 Vincenzellus ruficollis (Panzer, 1794) 5 Xestobium plumbeum (Illiger, 1801) 1 Xestobium rufovillosum (De Geer, 1774) 2 Xyleborinus saxesenii (Ratzeburg, 1837) 9 Xyleborus dispar (Fabricius, 1792) 6 Xyleborus dryographus (Ratzeburg, 1837) 10 Xyleborus monographus (Fabricius, 1792) 11 Xylotrechus antilope (Schönherr, 1817) 1 Xylotrechus arvicola (Olivier, 1795) 1

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Table S4

A check‐list of ants, bees and wasps.

Species Number of plots cinctellus (Spinola, 1808) 2 Ampulex fasciata Jurine, 1807 1 nigricornis (Curtis, 1826) 1 Ancistrocerus parietinus (Linnaeus, 1761) 1 Ancistrocerus trifasciatus (Müller, 1776) 1 Bombus humilis Illiger, 1806 2 Bombus hypnorum (Linnaeus, 1758) 1 Bombus lapidarius (Linnaeus, 1758) 1 Bombus pascuorum (Scopoli, 1763) 5 Bombus pratorum (Linnaeus, 1761) 2 Bombus ruderarius (Müller, 1776) 1 Camponotus fallax (Nylander, 1856) 1 Camponotus ligniperdus (Latreille, 1802) 5 Chelostoma campanularum (Kirby, 1802) 1 varus Lepeletier & Brullé, 1835 1 Crossocerus walkeri (Shuckard, 1837) 1 bifasciatus (Geoffroy, 1785) 2 (Magretti, 1886) 2 Discoelius dufourii Lepeletier, 1831 1 Dolichoderus quadripunctatus (Linnaeus, 1771) 7 Hylaeus angustatus (Schenck, 1861) 1 Hylaeus communis Nylander, 1852 1 Hylaeus confusus Nylander, 1852 3 Lasius brunneus (Latreille, 1798) 11 Lasius platythorax Seifert, 1991 5 Nitela borealis Valkeila, 1974 2 Passaloecus corniger Shuckard, 1837 1 Pemphredon lugens Dahlbom, 1842 1 Pemphredon lugubris Latreille, 1805 1 Pemphredon morio Van der Linden, 1829 1 Pemphredon rugifer Dahlbom, 1843 1 Temnothorax affinis (Mayr, 1855) 6

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