Department of Physics, Chemistry and Biology

Master Thesis

Comparison of saproxylic beetle assemblages on four different broad-leaved tree species in south-eastern Sweden

Helena Johansson

LiTH-IFM- Ex--11/2428--SE

Supervisor: Nicklas Jansson, Linköpings universitet Examiner: Anders Hargeby, Linköpings universitet

Department of Physics, Chemistry and Biology Linköpings universitet SE-581 83 Linköping, Sweden

Avdelning, Institution Datum Division, Department Date Avdelningen för biologi Instutitionen för fysik och mätteknik 2011-06-03

Språk Rapporttyp ISBN Language Report category LITH-IFM-A-EX--—11/2428—SE

______Svenska/Swedish Licentiatavhandling ISRN x Engelska/English x Examensarbete ______C-uppsats

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______Övrig rapport Serietitel och serienummer ISSN

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Handledare URL för elektronisk version Supervisor: Nicklas Jansson

Ort Location: Linköping

Titel Title: Comparison of saproxylic beetle assemblages on four different broad-leaved tree species in south- eastern Sweden

Författare Author: Helena Johansson

Sammanfattning Abstract: Old hollow trees have declined in Europe and many saproxylic (wood-dwelling) beetles dependent on them are threatened. Several studies have been done on old hollow oaks and they have been shown to harbour a species-rich saproxylic beetle fauna. However, other broad-leaved trees might also be important to consider as supporting habitats. The aim of this study was to investigate to what extent saproxylic beetles are tree genus specialists. Pitfall traps and window traps were used to compare the saproxylic beetle fauna in oak, ash, norway maple and small- leaved lime in an area dominated by old oaks. 5,501 specimens of saproxylic beetles were found, belonging to 239 species of which 27 species were red-listed. There were significant differences in the saproxylic species composition between the four tree species, but with large overlaps. The saproxylic species found in oak overlapped to 66 % with norway maple, to 67 % with ash and to 70 % with small-leaved lime. About one third of the species in this study seem to be tree genus specialists. The conclusion is that other broad-leaved trees are important to consider in models as supporting habitats for oaks. However, to be able to save the whole fauna of saproxylic beetles, trees of all different tree species are needed.

Nyckelord Keyword:

Conservation, coleoptera, beetles, saproxylic, broad-leaved trees, hollow, preferences Table of contents 1 Abstract ...... 2 2 Introduction ...... 2 2 Material and methods ...... 3 2.1. Study sites ...... 3 2.2 Beetle trapping ...... 4 2.3 Identification of saproxylic beetles ...... 5 2.4 Data analyses ...... 5 3 Results ...... 6 3.1 Obligate saproxylic beetles ...... 6 3.2 Facultative saproxylic beetles ...... 7 4 Discussion ...... 11 4.1 Conservation implications ...... 13 5 Acknowledgement ...... 13 6 References ...... 13 Appendix 1. The hollow trees ...... 16 Appendix 2. Saproxylic species table ...... 18

1 Abstract Old hollow trees have declined in Europe and many saproxylic (wood-dwelling) beetles dependent on them are threatened. Several studies have been done on old hollow oaks and they have been shown to harbour a species-rich saproxylic beetle fauna. However, other broad-leaved trees might also be important to consider as supporting habitats. The aim of this study was to investigate to what extent saproxylic beetles are tree genus specialists. Pitfall traps and window traps were used to compare the saproxylic beetle fauna in oak, ash, norway maple and small-leaved lime in an area dominated by old oaks. 5,501 specimens of saproxylic beetles were found, belonging to 239 species of which 27 species were red-listed. There were significant differences in the saproxylic species composition between the four tree species, but with large overlaps. The saproxylic species found in oak overlapped to 66 % with norway maple, to 67 % with ash and to 70 % with small-leaved lime. About one third of the species in this study seem to be tree genus specialists. The conclusion is that other broad-leaved trees are important to consider in models as supporting habitats for oaks. However, to be able to save the whole fauna of saproxylic beetles, trees of all different tree species are needed.

Keywords: Conservation, coleoptera, beetles, saproxylic, broad-leaved trees, hollow, preferences.

2 Introduction In Europe, many species dependent on old trees are threatened, due to habitat loss (Speight 1989). Old-growth temperate deciduous forests have decreased to a small fraction from their previous large distribution, thus old trees have disappeared from many regions (Hannah et al. 1995). Until the nineteenth century, old trees were common in pasture woodland and wooded meadows, but changes in land use have reduced the distribution of old trees (Kirby & Watkins 1998). When trees get older, structures serving as microhabitats for many organisms start to develop (Sverdrup-Thygeson et al. 2009). One of these microhabitats is trunk hollows that start to develop when the trees are about 200 years old (Ranius 2009a). Hollows in old trees provide important habitats for mammals, birds and many invertebrates. Tree hollows in the trunks often contain wood mould, which is loose wood colonized with fungi and remains from bird nests, insects and dead leaves (Ranius 2002a, Jansson et al. 2009a). Hollows with wood mould in old trees harbour a specialized insect fauna, among them saproxylic insects (Dajoz 2000). Saproxylic insects are insects that some part of their life cycle are dependent on dead wood (Speight 1989). Jonsell et al. (1998) suggest that the richness and composition of the red-listed saproxylic invertebrate species in boreal forests differ between different tree species. Many saproxylic insects were classified as specific to host tree genera and some were classified as monophagus to a single tree species. Results from field studies are mixed. A field study of saproxylic beetles on logging residues of aspen, birch, oak and spruce showed that the assemblages differed between tree species (Jonsell et al 2007) but Irmler et al (1996) found small differences between tree species on logs and stumps of three different tree species (beech, alder and spruce). Hollow oaks have been shown to harbour a rich saproxylic beetle fauna and it has been shown that different qualities of hollows and oak individuals affect the fauna (Ranius 2002b). However, there seem to be no systematic sampling comparing the saproxylic fauna in hollows between tree species.

2 The aim of this study was to investigate if different saproxylic beetles prefer hollow trees of different species. To what extents are saproxylic beetles tree genus specialists? A better knowledge about the habitat requirements of rare saproxylic beetles can be useful in landscape planning and conservation of this fauna. Old oaks have been in focus in many studies but other broad-leaved tree species might also be important to consider as supporting habitats. Therefore it is important to know which saproxylic beetles that utilize other broad-leaved tree genera and how specialized they are. This can be used to get better models in conservation and management planning.

2 Material and methods

2.1. Study sites This study was conducted in one of the few remaining landscapes in Northern Europe with high density of old oaks, in Östergötland, south-eastern Sweden (Antonsson & Wadstein 1991). Hollow trees of the different tree species, were selected from a database at the County Administration Board of Östergötland with old and hollow trees that they had registered the past 10 years was used. In a study area of approximately 10 x 30 km2, 48 hollow trees were selected, clustered at 10 study sites (Figure 1). At each site, at least one hollow oak (Quercus robur) was examined together with one or several hollow tree species of ash (Fraxinus excelsior), norway maple (Acer platanoides) and/or small-leaved lime (Tilia cordata) (Figure 2). The distance between two trees belonging to the same site was between 14-2603 m. In the study area with a buffer zone of 2603 m added to the rectangle, the percentage of old and hollow trees in the surrounding were approximately 68 % oaks, 8 % small-leaved lime, 6 % ash and 6 % norway maple according to the database at the County Administration Board.

Figure 1 Location of the 10 study sites in Östergötland, Sweden

3 6485000

6480000

6475000

6470000 Latitude 6465000

6460000

6455000 Quercus robur Fraxinus excelcior Acer platanoides Tilia cordata 6450000 1475000 1480000 1485000 1490000 1495000 1500000 1505000 1510000 Longitude

Figure 2 Spatial distribution of the 48 hollow trees, clustered in 10 study sites in Östergötland, Sweden (RT90)

2.2 Beetle trapping Beetles were collected using two kinds of traps – pitfall traps and window traps. The pitfall trap was a 7 cm ∅ plastic jar placed in the tree hollow with the opening level with the wood mould surface. The window trap was of a 30x50 cm2 wide, vertical transparent plastic plate with a tray underneath and was placed near (within 1 m) from the entrance of the tree hollow (Figure 2). The positions of the windows were 1.5-4 m above the ground, depending on where the hollow entrance was situated on the tree. The trays were filled to 70% with a solution of about 50% water, 50% propylene glycol, some denaturated alcohol to prevent animals from drinking the solution, and a few drops of detergent to reduce surface tension. The traps were in the trees from May to the later part of August and were emptied four times during that time. The target in this project was to have one oak with one or several of the other tree species per site, with one pitfall trap and one window trap on each tree. This target was not reached in some cases because the wood mould was too far away from the hollow entrance. Furthermore, 8 window traps were destroyed and therefore excluded from the study. In total, 48 trees were used in the study: 38 trees with pitfall trap and window trap, eight with only pitfall trap and two with only window trap (Table 1). According to a study comparing methods to survey saproxylic beetles (Ranius & Jansson 2002), window traps collect the highest number of species. Pitfall traps collect beetles associated with tree hollows, which is more rarely the case with

4 window traps. Therefore, to be able to catch a wide variety of saproxylic species, a combination of these two methods was chosen in this study.

Figure 2 The pitfall and window trap used in the study to collect saproxylic beetles

Table 1 Number of hollow tree species per site with the different traps

Site (oak coordinate) Acer Fraxinus Quercus Tilia Number of trees Number of trees Number of platanoides excelsior robur cordata with pitfall and with only pitfall trees with only window traps trap window trap

6478794, 1482072 3 1 1 2 7 6474590, 1481086 3 3 2 4 4 6473671, 1485330 2 1 2 3 1 1 6473712, 1489152 1 1 1 1 4 6471235, 1493164 3 1 1 4 1 6467243, 1496253 1 1 1 3 6465010, 1495177 2 1 1 1 5 6465498, 1492397 1 1 2 4 6461848, 1497758 2 2 1 1 2 6453697, 1491714 1 1 1 2 1

Total 13 11 12 12 38 8 3

2.3 Identification of saproxylic beetles Nicklas Jansson (NJ) identified most of the saproxylic beetles genera to species level, while other experts were consulted for the remaining genera (see Acknowledgement). The family Latrididae was left out from the study for financial reason. Species were classified as obligate or facultative saproxylic according to Dodelin et al. (2008), except for Alphitobius diaperinus, Antribus nebulosus, Korynetes caeruleus and Malachius bipustulatus that, based on own field experience by NJ, were classified as obligate saproxylic. Non-saproxylic species were left out from the study. Nomenclature follows Lundberg (1995) and the red-list categories Gärdenfors (2010).

2.4 Data analyses To compare the total species richness of saproxylic beetles of the different tree species, rarefaction curves with confidence interval and the expected number of species were calculated using the internet-accessible Species Richness Estimators Eco-Tool (Russell 2006), which implements the techniques described in Colwell and Coddington (1994) and references therein, as well as those of Colwell, Mao and

5 Chang (2004). The calculation of the expected numbers of species was done since the different tree species had different numbers of traps. Multivariate statistical analyses were made with the software CANOCO 4.5 (ter Braak and Smilauer 2002). The data was log transformed to minimize the impact of abundant species and rare species were down-weighted with the functions in Canoco (mainly because of a high percentage of singletons; Table 2). To evaluate the similarities of the saproxylic beetle fauna between the different tree species, a partial Detrended Correspondance Anlyses (pDCA) was made with site as a number of categorical covariables to eliminate potential spatial variation in the data. To test if the beetle fauna differed between the different tree species, a Monte Carlo permutation test with 9999 permutations was made (hence testing at α=0.0001) in a Canonical Correspondence Analysis (pCCA).

3 Results In total 5,501 individuals of saproxylic beetles were found in this study, belonging to 239 species. Of the total number of species, 27 were on the Swedish red-list (Gärdenfors 2010). The saproxylic species found in Quercus robur overlapped to 66 % with Acer platanoides, to 67 % with Fraxinus excelsior and to 70 % with Tilia cordata. A large proportion of the species were represented by only one individual. On the other hand, some saproxylic beetles were very abundant, e.g. Atheta nigricornis with 887 individuals.

3.1 Obligate saproxylic beetles There were small or no differences in the number of obligate saproxylic species found in pitfall traps and window traps between Acer platanoides, Fraxinus excelsior, Quercus robur and Tilia cordata (Table 2). The range of expected number of species overlap between the different tree species. The highest numbers of individuals and species were found in window traps. There were significant differences in the obligate saproxylic species composition both in pitfall traps and window traps between the four tree species (P = 0.0141; P = 0.001; Figure 3A, B). The obligate saproxylic fauna in pitfall traps in F. excelsior and T. cordata were more similar while the fauna of Q. robur and A. platanoides were more different. However, in window traps the obligate saproxylic fauna were more similar in F. excelsior and A. platanoides while the fauna of Q. robur and T. cordata were more different. There were obligate saproxylic species associated to all different tree species in both pitfall traps and window traps. In pitfall traps the species Euglenes oculatus, Dorcatoma chrysomelina, Dorcatoma flavicornis and Mycetophagus piceus showed strongest association to Q. robur. Phloeophagus lignarius and Korynetes caeruleus showed strongest association to A. platanoides. Prionychus ater, Melanotus castanipes and Gnathoncus spp showed strongest association to T. cordata. Many species had some association to F. excelsior, e.g. Diaperis boleti and Mycetochara axillaris. In window traps Dorcatoma chrysomelina, Dorcatoma flavicornis and Mycetophagus piceus showed association to Q. robur. Prionychus ater showed association to T. cordata. The same pattern as in the pitfall traps. Korynetes caeruleus showed association to T. cordata in the window traps instead of A. platanoides as in the pitfall traps. Even though there were significant differences between the faunas when analyzing the intermediate species composition of the different tree species, there was

6 large overlap between individual trees in the obligate saproxylic species composition caught by pitfall traps (Figure 4A). There were also large overlap between tree individuals in the tree species composition caught by window traps, but less compared to the pitfall traps (Figure 4B).

3.2 Facultative saproxylic beetles There were small or no differences found in the number of facultative saproxylic species found in pitfall traps and in window traps between Acer platanoides, Fraxinus excelsior, Quercus robur and Tilia cordata (Table 2). The range of expected number of species overlap between the different tree species. The highest numbers of individuals and species were found in window traps. There were significant differences in the facultative saproxylic species composition in pitfall traps between the four tree species (P = 0.0436; Figure 3C). However, there were no differences in the facultative species composition caught by window traps (graph not shown). The facultative saproxylic fauna in pitfall traps in A. platanoides and T. cordata were more similar while Q. robur and F. excelsior were more different. The species Palorus depressus showed strong association to Fraxinus excelsior and Cryptophagus dentatus showed strong association to Quercus robur. Even though there were significant differences between the faunas when analyzing the intermediate species composition of the different tree species, there was large overlap between individual trees in the facultative saproxylic species composition caught by pitfall traps (Figure 4C).

7 Table 2. Number of individuals and species of obligate and facultative saproxylic beetles divided in pitfall traps and window traps. CI in parenthesis. Acer Fraxinus Quercus Tilia Total Obligates Pitfall traps No. traps 13 11 11 10 34 No. ind. 144 175 231 84 634 No. species 39 (30-48) 44 (37-53) 47 (39-55) 35 (26-46) 93 Expected no. species 82 (55-157) 74(56-121) 81 (60-132) 114 (64-253) No. unique species 5 6 4 2 17 Singletons (%) 49 42 49 67 35 No. ind. red-listed 47 5 11 6 69 No. species red-listed 2 1 3 1 7 Window traps No. traps 10 9 11 10 40 No. ind. 496 441 1146 407 2490 No. species 80 (69-91) 78 (67-89) 105 (93-117) 79 (68-90) 115 Expected no. species 128 (102-187) 127 (101-183) 157 (130-214) 134 (104-202) No. unique species 5 6 8 2 21 Singletons (%) 74 77 57 61 40 No. red-listed ind. 21 8 63 18 110 No. red-listed species 8 7 15 9 24 Facultatives Pitfall traps No. traps 13 11 11 10 34 No. ind. 185 200 122 223 730 No. species 25 (19-31) 22 (16-28) 21 (14-28) 20 (15-25) 44 Expected no. species 41 (30-81) 39 (27-81) 45 (28-105) 32 (23-66) No. unique species 5 3 2 3 13 Singletons (%) 52 45 52 45 34 No. ind. red-listed 0 0 0 0 0 No. species red-listed 0 0 0 0 0 Window traps No. traps 10 9 11 10 40 No. ind. 509 245 370 523 1648 No. species 27 (23-33) 25 (19-31) 29 (24-34) 27 (21-33) 32 Expected no. species 35 (30-54) 38 (29-73) 37 (31-58) 39 (30-70) No. unique species 4 0 0 1 5 Singletons (%) 65 65 71 59 63 No. red-listed ind. 0 0 0 0 0 No. red-listed species 0 0 0 0 0

8

9

Figure 3. Ordination graphs showing species. The intermediate species composition of the saproxylic beetles in four different tree species (sample scores from a partial CCA). A) Obligate saproxylic beetles in pitfall traps. Species weight range 12%. B) Obligate saproxylic beetles in window traps. Species weight range 20%. C) Facultative saproxylic beetles in pitfall traps. Species weight range 1%.

10

Figure 4. Similarity (sample scores from a partial DCA) of the saproxylic beetle fauna in four different tree species. Each plot represents one tree individual. A) Obligate saproxylic species in pitfall traps. Eigenvalues for axis 1 and 2 were 0.689 and 0.419 respectively. B) Obligate saproxylic species in window traps. Eigenvalues for axis 1 and 2 were 0.286 and 0.201 respectively C) Facultative saproxylic species in pitfall traps. Eigenvalues for axis 1 and 2 were 0.318 and 0.228 respectively.

4 Discussion It has been concluded that the most important substrates for saproxylic invertebrates are veteran trees (trees with aging characteristics e.g. hollows) (Davies et al 2008). Quercus robur is considered to be the most species-rich tree (Jonsell et al 1998, Irmler el ta 2010) and the focus in several studies of saproxylic beetles, including the hermit beetle (Osmoderma eremita) (e.g. Ranius 2002a, Jansson et al 2009b). However, O. eremita has been shown to prefer Tilia cordata except for Q. robur and also to some small extent Franxinus excelsior and Acer platanoides in Poland (Oleksa et al. 2007). Jonsell et al (1998) made a study of substrate requirement including host specificity of all red-listed saproxylic beetles in Sweden based on faunistic literature and field experience. They found that almost 30% of the red-listed saproxylic invertebrates found in Q. robur were monophagus. However, systematic field studies of the importance of deciduous veteran trees of other tree species than Q. robur have been few including studies on tree host specificity of saproxylic beetles on veteran trees. In this study, large overlap was found in the saproxylic species composition between

11 hollow trees of Q. robur, A. platanoides, F. excelsior, and T. cordata. About two thirds of the saproxylic species found in Q. robur were found in A. platanoides, F. excelsior and T. cordata. Even though there were large overlaps in this study, there were significant differences in the saproxylic species composition between the four tree species. The obligate saproxylic fauna in pitfall traps in F. excelsior and T. cordata were more similar while the fauna of Q. robur and A. platanoides were more different. This is consistent with the similarity cluster analysis of the red-listed saproxylic invertebrates by Jonsell 1998. The obligate species, Dorcatoma chrysomelina, Dorcatoma flavicornis, Mycetophagus piceus and Euglenes oculatus were found strongest associated with Q. robur in this study. They are specialists on red-rotten oak wood and such wood is usually created through decay by the fungus Laetiporus sulphureus (Jansson et al 2009a). Dorcatoma flavicornis have been found on dead wood except for Q. robur, on A. platanoides by Jonsell 1998. In a study conducting saproxylic beetles on hollow T. cordata in Södermanland, Uppsala and Västmanland in Sweden, the species Dorcatoma chrysomelina, Dorcatoma flavicornis, Mycetophagus piceus and Euglenes oculatus were found on T. cordata (Jonsell & Sahlin 2010). In another study on islands dominating by T. cordata in Hjälmaren Sweden, only one individual of D. chrysomelina was found one time (Jonsell 2010). The other three species found strongest association to Q. robur in this study were absent. These mixed results show that the species were not host specific to Q. robur, but further studies are needed to be able to specify the levels of host specificity. Phloeophagus lignarius and Korynetes caeruleus were found strongest associated with A. platanoides in this study. P. lignarius live in hard white-root wood, which is most often found in A. platanoides and K. caeruleus is a predator on the species when larva. Prionychus ater, Melanotus castanipes and Gnathoncus spp showed strongest association to T. cordata. Prionychus ater live in the wood mould in most of tree species and M. castanipes and G. spp are predators, thus other species present decides their presence. Diaperis boleti and Mycetochara axillaris were found strongest associated to F. excelsior. D. boleti live on the fruiting body of the fungus Laetiporus sulphureus and M. axillaris on soft white-root wood. The tree species associations for the obligate saproxylic species in this study were consistent with the classifications by Jonsell 1998, except for Phloeophagus lignarius in pitfall traps and Anobium thomsoni and Xyleborinus saxeseni in window traps. In this study, the facultative species Palorus depressus showed strong association to F. excelsior but their presence depend on prey as most of the other species shown. Cryptophagus dentatus showed strong association to Quercus robur, which cannot be explained by previous knowledge. There seem to be no other systematic sampling comparing the saproxylic fauna in hollows between tree species, but there are some studies investigating tree host specificity on other dead wood with mixed results. Studies on high stumps (Lindhe & Lindelöw 2004) and on logging residues (Jonsell et al 2007) of spruce, birch, aspen and oak, found that the different tree species hosted different assemblages of saproxylic beetles. But between the deciduous trees, there was considerable overlap, especially between birch and oak (Lindhe & Lindelöw 2004). Another study on logs and stumps, found small differences between the tree species beech, alder and spruce (Irmler 1996). There also seem to be differences within the trees. The obligate saproxylic species composition inside the tree hollows (caught by pitfalls) was more similar between tree species than the species composition found outside the hollows (caught by window traps). Jonsell et al (1998) suggests that the wood of different tree species

12 become more similar when the wood decays and then tree species become less important. The substrate outside the tree hollows is probably more related to tree host- specific fungi and that may cause the larger differences in species composition for the species caught by window traps. There are also other factors than tree species that may affect the species composition. Larger trunks have been shown to harbor more saproxylic species because of higher amount of wood mould, which contributes to a more stable microclimate (Ranius & Jansson 2000). Sun exposure has also been shown to increase the species richness of saproxylic beetles. The preference of sun-exposed wood has been stated in several studies (Gärdenfors & Baranowski 1992, Jonsell et al 1998, Alexander 1999, Lindhe 2005). Most trees in this study were situated in sun-exposed avenues. This study was conducted in a landscape with high density of old oaks (Antonsson & Wadstein 1991). About 70 % of the old hollow trees in the study area were Q. robur and Q. robur was also the most species-rich species in the study. In line with this, Fagus sylvatica had higher number of saproxylic beetle species than oaks in a study in a F. sylvatica dominated area in Germany (Irmler et al 2010). The oaks in the surrounding might act as sources and the other tree species as sinks. Without Q. robur as sources in the surrounding area, the species are absent in the other broad- leaved tree species.

4.1 Conservation implications Quercus robur is considered to be the most species-rich tree genera and generally species-rich tree genera also have the largest numbers of specific species (Jonsell et al 1998). However, in this study all the four tree species had some monophagus (specific) species associated. About one third of the species in this study seem to be tree genus specialists. Other broad-leaved trees are important to consider in models as supporting habitats for oaks. However, the other broad-leaved tree species decay faster than the oak and therefore serves as support during a shorter time. Though, an older hollow broad-leaved tree should be considered more attention for saving than young oaks. To be able to save the whole fauna of saproxylic beetles, trees of all different tree species are needed.

5 Acknowledgement I would like to thank my supervisors Nicklas Jansson, Karl-Olof Bergman and Per Milberg for their support during this project and comments on this manuscript. The landowners for their permission to put up traps on their trees and the entomologists Stig Lundberg and Rickard Andersson for identification of some of the species. Financial support to this project came from Eklandskapsfonden the municipality of Linköping and The County Administration board of Östergötland.

6 References Antonsson, K., Wadstein, M., 1991. Eklandskapet. En naturinventering av hagar och lövskogar I Eklandskapet söder om Linköping. Länsstyrelsen I Östergötlands län, Linköping (in Swedish) Alexander, K., 1999. Should deadwood be left in sun or shade?. British Wildlife, 10: 342

13 Davies, Z.G., Tyler, C., Stewart, G.B., Pullin, A.S., 2008. Are current management recommendations for saproxylic invertebrates effective? A systematic review. Biodiversity and Conservation, 17: 209-234 Dajoz, R., 2000. Insects and forests. The role and diversity of insects in the forest environment. Intercept, Londres Dodelin, B., Ballot, P., Stokland, J.N., 2008. The saproxylic database checklist of beetles vers. 8/2008: www.saproxylic.org Cited in Dec 2010. Gärdenfors, U., (Ed.) 2010. Rödlistade arter i Sverige 2010—The 2010 red list of Swedish species. Swedish Species Information Centre, SLU, Uppsala Gärdenfors, U., Baranowski, R., 1992. Skalbaggar anpassade till öppna respektive slutna ädellövskogar föredrar olika trädslag (Beetles living in open forests prefer different tree species than those living in dense forests). Entomologisk Tidskrift, 113: 1-11 (in Swedish, English abstract) Hannah, L., Carr, J.L., Lankerani, A., 1995. Human disturbance and natural habitat: a biome level analysis of a global data set. Biodiversity and Conservation, 4: 128- 155 Irmler, U., Heller, K., Warning J., 1996. Age and tree species as factors influencing the populations of insects living in dead wood (Coleoptera, Diptera: Sciaridae, Mycetophilidae). Pedobiologia, 40: 134-148 Irmler, U., Arp, H., Nötzold, R., 2010. Species richness of saproxylic beetles in woodlands is affected by dispersion ability of species, age and stand size. Journal of Insect Conservation, 14: 227-235 Jansson, N., Ranius, T., Larsson, A., Milberg, P., 2009a. Boxes mimicking tree hollows can help conservation of saproxylic beetles. Biodiversity and Conservation, 18: 3891-3908 Jansson, N., Bergman, K-O., Jonsell, M., Milberg, P., 2009b. An indicator system for identification of sites of high conservation value for saproxylic oak (Quercus spp.) beetles in southern Sweden. Journal of Insect Conservation, 14: 399-412 Jonsell, M., Westlien, J., Ehnström, B., 1998. Substrate requirements of red-listed saproxylic invertebrates in Sweden. Biodiversity and Conservation, 7: 749-764 Jonsell, M., Hansson, J., Wedmo L., 2007. Diversity of saproxylic beetle species in logging residues in Sweden – Comparisons between tree species and diameters. Biological and Conservation, 138: 89-99 Jonsell, M., Sahlin, E., 2010. Inventering av vedlevande skalbaggar på lindar i Södermanlands, Uppsala och Västmanlands län. Jonsell, M. 2010. Inventering av vedlevande skalbaggar – i lindmiljöer på öar i Hjälmaren. Länsstyrelsen i Örebro län, publ.nr. 2010:1. In Swedish. Kirby, K.J, Watkins, C., 1998. The ecological history of European forests. CAP International, Oxon Lindhe, A., Lindhelöw, A., 2004. Cut high stumps of spruce, birch, aspen and oak as breeding substrates for saproxylic beetles. Forest Ecology and Management. 203: 1-20 Lindhe, A., Lindelöw, Å., Åsenblad N., 2005. Saproxylic beetles in standing dead wood density in relation to substrate sun-exposure and diameter. Lundberg, S., 1995. Catalagus Coleopterorum Sueciae. Naturhistoriska Riksmuseet. Stockholm, Sweden Palm, T., 1959. Die Holz- und Rindenkäfer der Süd- und Mittelschwedischen Laubbäume (The wood and bark coleoptera of deciduous trees in south and central Sweden). Opuscula Entomologiska Supplementum XVI. (in German, English summary).

14 Ranius, T., 2002a. Influence of stand size and quality of tree hollows on saproxylic beetles in Sweden. Biological Conservation, 103: 85-91 Ranius, T., 2002b. Population ecology and conservation of beetles and pseudoscorpions living in hollow oaks in Sweden. Animal Biodiversity and Conservation, 25: 53-68 Ranius, T., Jansson, N., 2002. A comparison of the three methods to survey saproxylic beetles in hollow oaks Ranius, T., Niklasson, M., Berg, N. 2009. Development of tree hollows in pedunculate oak (Quercus robur). Forest Ecology and Management, 257: 303-130 Russell, G. J., 2006. Species Richness v2.1. Eco-Tools web site: http://www.eco- tools.net Speight, M.C.D., 1989. Saproxylic invertebrates and their conservation. Council of Europe, Strasbourg. Sverdrup-Thygeson, A., Skarpaas O., Ödegaard, F., 2009. Hollow oaks and beetle conservation: the significance of the surroundings. Biodiversity and Conservation, 19: 837-852 Tavakilian, G., Berkov, A. and Meuer-Grimes, B., 1997. Neotrophical tree species and their faunas of Xylophagus Longicorns (Coleoptera: Cerambycide) in French Guiana. The Botanical Review, 63, 303-305 ter Braak, C.J.F., Smilauer, P., 2002. Canoco reference manual and user’s guide to Canoco for windows: software for Canonical Community Ordination (version 4). Microcomputer Power, Ithaca

15 Appendix 1. The hollow trees PT = Pitfall trap WT = Window trap

Area Tree species Nr RT90 X RT90 Y Traps Comment Bjärka-Säby Quercus robur 1 6461848 1497758 PT+WT Bjärka-Säby Quercus robur 2 6461852 1497745 WT WT destroyed during a period and excluded in the results. Bjärka-Säby Tilia cordata 1 6461646 1497874 PT+WT Bjärka-Säby Tilia cordata 2 6461822 1497668 WT Wood mould too far away for PT Brokind Acer platanoides 6454240 1491358 PT Brokind Quercus robur 6453697 1491714 PT+WT Brokind Tilia cordata 6455179 1491149 PT+WT Gerstorp Acer platanoides 1 6478656 1482454 PT+WT Gerstorp Acer platanoides 2 6478582 1482468 PT+WT Gerstorp Acer platanoides 3 6478549 1482474 PT+WT Gerstorp Fraxinus excelsior 6478978 1482991 PT+WT Gerstorp Quercus robur 6478794 1482072 PT+WT Gerstorp Tilia cordata 1 6478508 1481937 PT+WT Gerstorp Tilia cordata 2 6478633 1482382 PT+WT Harvestad Fraxinus excelsior 1 6470943 1493407 PT+WT Harvestad Fraxinus excelsior 2 6470733 1493540 PT+WT Harvestad Fraxinus excelsior 3 6472406 1491830 PT+WT Harvestad Quercus robur 6471235 1493164 PT+WT Harvestad Tilia cordata 6470880 1493939 PT+WT WT destroyed during a period and excluded in the results. Hovetorp Acer platanoides 1 6464455 1495542 PT+WT Hovetorp Acer platanoides 2 6463766 1495443 PT+WT Hovetorp Fraxinus excelsior 6464627 1495405 PT+WT Hovetorp Quercus robur 6465010 1495177 PT+WT Hovetorp Tilia cordata 6465301 1494588 PT+WT Lagerlunda Acer platanoides 1 6474095 1481972 PT+WT WT destroyed during a period and excluded in the results. Lagerlunda Acer platanoides 2 6474180 1481987 PT+WT Lagerlunda Acer platanoides 3 6474394 1481427 PT+WT WT destroyed during a period and excluded in the results. Lagerlunda Fraxinus excelsior 1 6474629 1481820 PT+WT WT destroyed during a period and excluded in the results. Lagerlunda Fraxinus excelsior 2 6474394 1481405 PT+WT WT destroyed during a period and excluded in the results. Lagerlunda Fraxinus excelsior 3 6474234 1481143 PT+WT Lagerlunda Quercus robur 1 6474590 1481086 PT+WT Lagerlunda Quercus robur 2 6474630 1481065 PT+WT

16 Lambohov Acer platanoides 1 6472820 1485520 PT+WT Lambohov Acer platanoides 2 6472618 1485374 PT+WT Lambohov Quercus robur 6473671 1485330 PT+WT Lambohov Tilia cordata 1 6472861 1485590 PT+WT WT destroyed during a period and excluded in the results. Lambohov Tilia cordata 2 6472665 1485538 PT+WT PT generated extreme values and excluded in the results. Smedstad Acer platanoides 6473533 1488969 PT+WT Smedstad Fraxinus excelsior 6473709 1489166 PT+WT Smedstad Quercus robur 6473712 1489152 PT+WT Smedstad Tilia cordata 6473681 1489182 PT+WT Stavsätter Fraxinus excelsior 6464801 1491918 PT+WT Stavsätter Quercus robur 6465498 1492397 PT+WT Stavsätter Tilia cordata 1 6463533 1492309 PT+WT Stavsätter Tilia cordata 2 6465684 1492211 PT+WT Sturefors Acer platanoides 6466959 1496975 PT+WT Sturefors Fraxinus excelsior 6467055 1496525 PT+WT Sturefors Quercus robur 6467243 1496253 PT+WT

17 Appendix 2. Saproxylic species table PT = Pitfall trap WT = Window trap

T

W

PT

Cat Nr Species RL Acer No. Ind. PT Fraxinus No. Ind. PT Quercus No. Ind Tilia No. Ind. PT Acer No. Ind. WT Fraxinus No. Ind. Quercus No. Ind. WT Tilia No. Ind. WT

Histeridae

O 652 Plegaderus caesus 0 1 0 0 0 1 1 0 F 672 Gnathoncus nannetensis 0 0 0 0 0 0 1 0 O 673 Gnathoncus spp 2 4 1 5 3 0 3 5 F 674 Gnathoncus buyssoni 1 3 0 0 1 0 2 0 F 677 Dendrophilus corticalis 17 74 7 66 0 1 0 3 O 680 Paromalus flavicornis 1 14 6 0 0 0 5 0 Leioddidae O 842 Anisotoma humeralis 0 0 0 0 0 1 4 0 O 845 Anisotoma glabra 0 0 0 0 1 0 1 0 O 863 Agathidium nigripenne 0 0 0 0 2 0 0 0 O 865 Agathidium seminulum 0 0 0 0 0 0 0 2 O 868 Agathidium pisanum 0 0 1 0 0 0 0 0 O 888 Nemadus colonoides 0 1 0 0 1 0 0 0 Scydmaenidae O 946 Stenichnus scutellaris 0 1 0 2 0 0 0 1 O 948 Stenichnus godarti 0 0 0 0 0 2 0 1 Staphylinidae F 994 Gabrius splendidulus 0 0 0 0 0 0 1 0 F 1101 Velleius dilatatus 0 1 0 1 0 2 0 1 F 1105 Quedius mesomelinus 74 30 63 54 70 43 70 92 F 1107 Quedius cruentus 5 4 1 0 21 3 5 7 F 1108 Quedius invreai 0 0 0 0 0 0 0 1 O 1109 Quedius brevicornis 0 4 0 1 0 0 1 0 O 1111 Quedius brevis 0 0 1 1 0 1 1 0 O 1113 Quedius microps 1 0 1 3 0 0 0 3 F 1118 Quedius xanthopus 0 0 1 1 2 1 0 1 O 1162 Nudobius lentus 0 0 0 0 0 0 1 0 F 1170 Xantholinus tricolor 1 0 0 0 0 0 0 0 O 1331 Bibloporus minutus 0 0 0 0 0 0 0 1 O 1350 Euplectus fauveli 0 0 0 0 0 0 1 0 O 1368 Trichonyx sulcicollis NT 0 0 0 0 1 1 0 0 F 1387 Megarthrus depressus 1 1 0 0 0 0 1 0 F 1412 Phyllodrepa melanocephala 1 0 0 0 1 0 0 2 F 1414 Phyllodrepa nigra 0 1 0 0 0 0 0 4

18 O 1419 Phyllodrepa linearis 0 0 0 0 0 0 1 0 O 1421 Phyllodrepa ioptera 0 0 0 0 1 0 0 0 O 1424 Hapalaraea pygmaea 1 0 0 0 0 0 0 0 O 1446 Phloeonomus sjobergi 0 0 0 0 1 0 0 0 F 1497 Scaphisoma boleti 3 0 0 0 0 0 0 0 O 1501 Scaphisoma assimile 0 0 0 0 0 0 0 1 F 1504 Coprophilus striatulus 1 0 0 0 0 0 0 0 F 1538 Oxytelus laqueatus 0 0 0 0 2 1 0 1 O 1539 Anotylus insecatus 0 0 0 0 0 0 1 0 O 1623 Carphacis striatus VU 0 0 0 0 0 0 1 1 F 1634 Sepedophilus littoreus 0 0 0 0 0 0 1 0 F 1635 Sepedophilus testaceus 5 0 1 1 0 0 0 0 F 1637 Sepedophilus constans 1 0 0 1 1 0 0 0 F 1639 Sepedophilus immaculatus 0 0 0 0 0 0 1 0 O 1688 Aleochara sparsa 6 1 4 1 20 17 58 27 F 1735 Oxypoda alternans 0 0 0 1 0 0 0 0 O 1737 Oxypoda arborea 0 0 0 2 0 0 0 0 F 1781 Haploglossa gentilis 4 10 6 47 1 1 8 6 F 1782 Haploglossa villosula 4 2 2 4 39 17 54 80 F 1785 Haploglossa marginalis 0 0 0 0 1 2 5 0 O 1800 Phloeopora corticalis 0 0 0 0 0 2 2 0 F 1925 Atheta sodalis 1 0 1 0 0 0 3 0 F 1926 Atheta gagatina 0 0 0 0 0 0 1 0 O 1969 Atheta castanoptera 0 0 0 0 2 1 0 0 O 1993 Atheta britanniae 3 0 0 0 7 8 1 2 F 1994 Atheta crassicornis 10 1 1 3 2 3 1 3 F 1998 Atheta euryoptera 0 0 0 0 0 0 1 0 F 2001 Atheta nigricornis 33 52 17 20 293 110 114 246 F 2002 Atheta harwoodi 0 1 0 0 0 0 0 0 O 2054 Thamiaraea cinnamomea 0 0 1 0 0 2 26 3 F 2068 Zyras funestus 0 0 0 12 0 0 0 0 F 2071 Zyras lugens 0 0 1 1 0 0 0 1 F 2072 Zyras laticollis 0 0 1 2 0 0 0 0 O 2120 Anomognathus cuspidatus 0 0 0 0 0 0 1 0 O 2128 Placusa depressa 0 0 0 0 1 3 1 2 O 2133 Placusa atrata 0 0 0 0 1 0 0 0 F 2144 Holobus flavicornis 0 0 0 0 1 0 0 0 Scirtidae O 2197 Prionocyphon serricornis 0 0 0 0 0 1 0 1 Scarabaidae F 2203 Trox scaber 1 2 0 1 1 1 1 0 F 2290 Cetonia aurata 0 1 0 0 0 3 2 0 O 2291 Liocola marmorata 0 2 0 0 0 6 0 2 O 2293 Osmoderma eremita NT 0 0 2 1 0 0 0 0 Lucanidae

19 O 2300 Platycerus caraboides 0 0 0 0 0 3 0 0 O 2302 Sinodendron cylindricum 2 3 4 1 1 0 2 1 Elateridae O 2410 Denticollis linearis 0 0 1 0 1 1 0 8 O 2423 Prosternon tessellatum 0 0 0 0 0 1 1 0 O 2432 Calambus bipustulatus NT 0 0 0 1 0 1 2 1 O 2434 Procraerus tibialis NT 1 0 1 0 0 0 12 0 O 2441 Ampedus nigroflavus NT 0 1 0 1 3 0 2 0 O 2442 Ampedus pomorum 0 1 0 0 0 0 2 1 O 2443 Ampedus hjorti 0 0 1 1 0 0 7 3 O 2447 Ampedus balteatus 0 0 4 0 0 1 4 0 O 2449 Ampedus praeustus NT 0 0 0 0 0 0 1 0 O 2450 Ampedus cardinalis NT 0 0 3 0 0 0 4 0 O 2456 Elater ferrugineus VU 2 0 0 0 0 0 0 0 O 2458 Melanotus villosus 1 0 0 0 0 7 4 0 O 2459 Melanotus castanipes 1 2 3 14 4 3 5 3 Eucnemidae O 2478 Microrhagus lepidus NT 0 0 0 0 0 0 0 1 O 2485 Hylis foveicollis 0 0 0 0 0 0 0 2 Buprestidae O 2514 Agrilus biguttatus NT 0 0 0 0 0 0 3 0 O 2519 Agrilus sulcicollis 0 0 0 0 0 0 1 0 Dermestidae F 2562 Dermestes lardarius 1 0 0 0 0 0 0 0 F 2566 Attagenus pellio 1 0 0 0 2 2 2 3 O 2578 Globicornis rufitarsis 0 0 0 0 1 0 10 1 F 2579 Megatoma undata 0 1 1 0 4 1 4 6 O 2581 Ctesias serra 0 2 0 0 23 9 56 21 F 2583 Anthrenus scrophulariae 3 0 0 0 1 0 6 0 O 2585 Anthrenus museorum 0 0 0 0 2 6 8 2 Anobiidae O 2617 Ptinus rufipes 2 1 0 1 7 1 29 10 F 2619 Ptinus fur 0 1 0 0 0 0 0 0 O 2625 Hedobia imperalis 0 1 0 0 1 0 1 1 O 2628 Xestobium rufovillosum 10 12 51 4 4 0 5 2 O 2640 Gastrallus immarginatus 0 0 0 0 0 0 5 0 O 2642 Anobium nitidum 12 18 0 11 30 22 23 19 O 2645 Anobium rufipes 1 0 4 0 2 0 2 1 O 2647 Microbregma emarginata 0 0 0 0 0 12 0 0 O 2648 Hadrobregmus pertinax 0 0 0 0 2 0 0 21 O 2651 Ptilinus pectinicornis 0 0 0 0 1 0 0 0 O 2652 Ptilinus fuscus 0 0 0 0 3 0 0 0 O 2656 Xyletinus pectinatus 0 0 0 0 0 0 4 0 O 2660 Xyletinus longitarsis VU 0 0 0 0 0 0 1 0 O 2665 Dorcatoma flavicornis 0 0 4 0 0 1 64 0

20 O 2667 Dorcatoma chrysomelina 0 6 27 0 1 19 172 1 O 2670 Dorcatoma dresdensis 0 0 0 0 1 0 0 0 O 2671 Dorcatoma robusta 0 0 0 0 1 0 1 0 Lymexylonidae O 2674 Hylecoetus dermestoides 0 0 0 0 4 9 0 0 O 2675 Lymexylon navale NT 0 0 0 0 0 0 1 0 Trogositidae O 2682 Grynocharis oblonga 5 10 7 3 1 0 6 0 Cleridae O 2687 Tillus elongatus 4 0 0 0 3 0 3 1 O* 2695 Korynetes caeruleus 9 4 4 0 5 3 10 16 Melyridae O 2703 Trichoceble floralis NT 0 1 0 1 1 0 2 1 O 2707 Dasytes niger 0 0 0 0 0 0 3 0 O 2709 Dasytes cyaneus 1 0 0 0 4 3 2 0 O 2712 Dasytes plumbeus 0 1 0 0 5 23 12 17 O 2716 Hypebaeus flavipes 0 0 0 0 0 0 3 0 O* 2725 Malachius bipustulatus 0 0 0 0 1 1 1 1 O 2729 Anthocomus fasciatus 0 0 0 0 0 1 0 0 Nitidulidae F 2742 Carpophilus marginellus 1 0 0 0 1 0 0 0 F 2745 Epuraea melanocephala 0 0 0 0 4 0 0 0 O 2746 Epuraea guttata 0 0 1 0 1 1 26 0 O 2759 Epuraea marseuli 0 0 0 0 0 0 0 3 O 2760 Epuraea pygmaea 0 1 0 0 0 0 0 0 F 2763 Epuraea placida 0 0 1 0 0 3 0 0 F 2764 Epuraea terminalis 0 0 0 1 0 0 0 1 F 2775 Epuraea melina 0 0 0 0 0 0 0 1 O 2826 Soronia grisea 0 1 0 0 35 32 30 22 F 2828 Pocadius ferrugineus 0 1 0 0 0 1 0 1 O 2834 Cryptarcha strigata 0 5 21 5 17 22 165 50 O 2835 Cryptarcha undata 0 0 0 0 3 5 26 4 F 2837 Glischrochilus hortensis 8 7 2 2 28 20 4 17 Monotomidae O 2847 Rhizophagus ferrugineus 0 0 0 0 0 1 0 0 F 2851 Rhizophagus dispar 1 0 0 2 3 1 0 2 O 2852 Rhizophagus bipustulatus 4 6 1 1 2 4 5 6 O 2853 Rhizophagus nitidilus 0 0 0 0 0 0 0 1 O 2856 Rhizophagus cribratus 0 0 1 0 0 1 0 2 Laemophloidae F 2887 Cryptolestes ferrugineus 0 0 2 0 0 0 1 0 Cryptophagidae O 2912 Cryptophagus badius 1 0 1 0 0 0 0 0 F 2913 Cryptophagus populi 0 0 0 0 0 0 0 1 O 2922 Cryptophagus micaceus 0 2 3 0 14 5 9 16

21 O 2923 Cryptophagus saginatus 0 0 1 0 0 0 1 0 O 2924 Cryptophagus fuscicornis VU 0 0 1 0 0 0 0 2 O 2926 Cryptophagus confusus 0 1 0 0 0 2 1 0 F 2928 Cryptophagus dentatus 0 0 6 2 3 3 11 11 Cryptophagus O 2929 pseudodentatus 0 0 0 0 1 1 4 2 F 2931 Cryptophagus distinguendus 0 0 1 0 0 0 0 0 O 2933 Cryptophagus scanicus 9 3 6 1 3 1 6 4 F 2937 Cryptophagus scutellatus 0 2 1 0 0 0 0 0 O 2939 Cryptophagus pilosus 1 0 1 0 0 0 0 1 O 2953 Atomaria morio 2 2 1 1 0 0 1 2 F 2962 Atomaria fuscata 0 0 0 0 2 1 0 0 O 2973 Atomaria nitidula 0 0 0 0 0 0 0 1 O 2983 Atomaria umbrina 0 0 0 1 0 0 0 0 O 2991 Atomaria nigriostris 0 0 0 0 0 1 0 0 O 3005 Atomaria procerula 1 0 0 0 0 0 0 0 Erotylidae O 3009 Tritoma bipustulata 0 0 0 0 0 0 0 1 O 3011 Triplax russica 4 0 0 0 3 1 2 0 O 3013 Triplax rufipes NT 0 0 0 0 1 0 0 0 O 3015 Dacne bipustulata 0 0 1 2 15 9 93 1 Cerylonidae O 3037 Cerylon histeroides 2 3 2 0 2 0 1 7 O 3038 Cerylon ferrugineum 0 1 1 1 0 1 7 0 Endomychidae F 3047 Mycetaea subterranea 0 0 0 0 0 0 1 0 O 3048 Leiestes seminigra NT 1 0 0 0 0 0 0 0 O 3052 Endomychus coccineus 0 0 0 0 1 0 0 0 Cisidae O 3213 Cis lineatocribratus 0 1 0 0 1 0 0 0 O 3214 Cis alter 0 1 0 1 0 0 0 0 O 3218 Cis comptus 0 0 0 0 0 0 2 0 O 3219 Cis hispidus 0 0 1 0 0 0 1 0 O 3223 Cis rugulosus NT 0 0 0 0 1 0 0 0 O 3229 Cis bidentatus 2 0 1 0 0 0 0 3 O 3230 Ennearthron cornutum 1 0 0 0 0 0 2 0 O 3234 Orthocis alni 0 0 0 0 1 0 0 0 Colydidae O 3249 Synchita humeralis 0 0 0 0 0 0 1 0 Mycetophagidae O 3257 Litargus connexus 0 0 0 0 2 1 1 1 O 3260 Mycetophagus piceus 0 1 10 0 6 16 75 4 Mycetophagus O 3265 multipunctatus 0 2 0 0 0 1 1 0 O 3266 Mycetophagus fulvicollis NT 0 0 0 0 0 0 2 2 O 3267 Mycetophagus populi 1 0 0 0 1 0 0 1

22 Oedemeridae O 3281 Ischnomera caerulea 1 0 0 0 2 0 0 0 O 3283 Ischnomera cinerascens NT 0 0 0 0 1 1 0 0 Pyrochroidae O 3295 Schizotus pectinicornis 0 0 0 0 1 0 1 0 Salpingidae O 3298 Lissodema cursor 0 0 0 0 0 1 0 0 O 3307 Salpingus planirostris 0 0 0 0 2 1 0 0 O 3308 Salpingus ruficollis 0 0 0 0 6 2 1 3 Aderidae O 3311 Pseudeuglenes pentatomus VU 0 0 0 0 0 0 0 1 O 3313 Euglenes pygmaeus 0 0 2 0 0 1 2 0 O 3314 Euglenes oculatus 0 0 32 0 0 2 26 0 Tenebrionidae O 3342 Eledona agaricola 0 9 4 0 0 0 1 0 O 3343 Diaperis boleti 1 3 2 1 16 10 22 8 F 3364 Palorus depressus 0 2 2 0 1 1 2 1 O* 3370 Alphitobius diaperinus 0 9 0 1 0 0 0 0 O 3378 Uloma culinaris NT 0 2 1 0 0 1 0 0 O 3383 Tenebrio opacus VU 0 0 1 0 0 0 0 0 F 3385 Tenebrio molitor 1 2 0 0 0 0 0 0 O 3395 Corticeus fasciatus VU 0 0 0 0 0 0 1 0 O 3397 Allecula morio NT 0 0 2 0 0 1 26 8 O 3400 Prionychus ater 0 11 3 8 0 7 6 10 O 3403 Pseudocistela ceramboides 1 2 0 0 0 2 5 23 O 3406 Mycetochara flavipes 0 1 0 0 1 0 1 2 O 3407 Mycetochara axillaris 4 17 0 1 89 46 4 4 O 3408 Mycetochara humeralis NT 0 0 0 0 7 2 2 0 O 3410 Mycetochara linearis 0 0 0 0 40 15 9 12 Scraptidae O 3414 Scraptia fuscula 0 0 1 1 0 8 6 6 O 3417 Anaspis frontalis 0 0 0 0 0 0 0 1 O 3419 Anaspis marginicollis 0 0 0 0 5 5 3 2 O 3420 Anaspis thoracica 0 0 2 1 3 4 10 2 O 3424 Anaspis rufilabris 0 0 0 1 3 3 2 6 Mordellidae O 3427 Tomoxia bucephala 1 0 0 0 2 9 3 3 O 3448 Mordellistena variegata 0 0 0 0 0 1 0 0 Melandryidae O 3459 Orchesia micans 0 0 0 0 0 0 1 0 O 3463 Orchesia undulata 0 0 0 0 2 3 3 4 Cerambycidae O 3498 Rhagium mordax 0 0 0 0 1 0 1 0 O 3500 Oxymirus cursor 0 0 0 0 1 0 0 0 O 3501 Stenocorus meridianus 0 0 0 0 0 1 1 0

23 O 3513 Grammoptera ruficornis 0 0 0 0 0 0 0 1 O 3514 Alosterna tabacicolor 0 0 0 0 2 0 1 7 Staphylinidae O 3556 Phymatodes testaceus 0 0 0 0 1 5 0 1 O 3556 Phymatodes testaceus 2 0 0 0 2 0 2 0 Cerambycidae O 3585 Leiopus nebulosus 0 0 0 0 0 1 0 0 O 3590 Exocentrus lusitanus 0 0 0 0 0 0 0 1 Anthribidae O* 3818 Anthribus nebulosus 0 0 0 0 2 0 0 0 Curculionidae O 4296 Rhyncolus ater 2 0 0 1 0 0 0 0 O 4299 Phloeophagus lignarius NT 43 1 0 2 6 1 0 1 O 4454 Hylesinus crenatus 0 0 0 0 0 4 1 0 O 4456 Hylesinus fraxini 0 0 0 0 1 23 0 0 O 4477 Scolytus intricatus 0 0 0 0 0 1 1 0 O 4500 Dryocoetes villosus 0 0 0 1 4 3 16 5 O 4515 Xyleborus monographus 0 0 0 0 1 0 0 0 O 4516 Xyleborinus saxesenii 1 0 0 0 56 23 9 4 * Classified as obligate saproxylic based on own field experience by Nicklas Jansson

24