445 The preferences of saproxylic species for different dead wood types created in forest restoration treatments

Tero Toivanen and Janne S. Kotiaho

Abstract: Restoration by imitating natural disturbances is widely practised in boreal forests to increase the availability of habitats for specialized species. We studied the abundance and species richness of saproxylic on different types of created dead wood during 2 years after restoration. The study was conducted on areas of a large-scale experiment in which Norway spruce ( (L.) Karst.) forests were restored by controlled burning and partial harvesting with down wood retention in southern . More beetle species were attracted to spruces than to birches and more species were attracted to burnt trees than to unburnt trees killed by girdling. Birch-living species consistently benefited from fire, but on spruce, the abundance of cambium consumers and their associates was negatively affected by fire. Trees at harvested sites attracted more beetles in the first year, but the volume of down wood retention had only minor effects. Beetle assemblages were strongly altered by burning and harvesting. We conclude that burning and harvesting are efficient tools to promote species richness within a short time period, but there is a risk that the dead wood resource may be rapidly exhausted. Moreover, many saproxylic species of spruce forests may not be adapted to open habitats formed by stand-replacing distur- bances. Re´sume´ : La restauration qui imite les perturbations naturelles est largement utilise´e dans les foreˆts bore´ales afin d’ac- croıˆtre la disponibilite´ des habitats pour les espe`ces spe´cialise´es. Nous avons e´tudie´ l’abondance et la richesse en espe`ces des cole´opte`res saproxyliques pendant 2 ans sur les diffe´rents types de bois mort apparu a` la suite d’une restauration. L’e´tude a e´te´ re´alise´e sur le territoire d’une expe´rience a` grande e´chelle ou` des foreˆts d’e´pice´a commun (Picea abies (L.) Karst.) ont e´te´ restaure´es par un bruˆlage dirige´ et une coupe partielle avec re´tention du bois au sol dans le sud de la Fin- lande. Plus d’espe`ces de cole´opte`res ont e´te´ attire´es par les e´pice´as que par les bouleaux et plus d’espe`ces ont e´te´ attire´es par les arbres bruˆle´s que par les arbres non bruˆle´s tue´s par anne´lation. Les espe`ces vivant sur le bouleau ont invariable- ment be´ne´ficie´ du feu. Par contre sur l’e´pice´a, l’abondance des cole´opte`res qui attaquent le cambium et leurs associe´s e´taient ne´gativement affecte´s par le feu. Les arbres sur les stations coupe´es ont attire´ plus de cole´opte`res durant la pre- mie`re anne´e mais le volume de re´tention de bois au sol a eu seulement des effets mineurs. Les assemblages de cole´opte`res ont e´te´ fortement modifie´s par le bruˆlage et la coupe. Nous concluons que le bruˆlage et la coupe sont des outils efficaces pour favoriser la richesse en espe`ces sur une courte pe´riode de temps mais qu’il y a un risque que les ressources en bois mort soient rapidement e´puise´es. De plus, plusieurs espe`ces saproxyliques associe´es aux foreˆts d’e´pice´a ne sont peut-eˆtre pas adapte´es aux habitats ouverts cre´e´s par les perturbations qui provoquent le remplacement des peuplements. [Traduit par la Re´daction]

Introduction dynamics of natural forests, such as reintroducing fire as a disturbance factor, to create structural elements and resour- To protect biodiversity in boreal forests, the need for ac- ces important for biodiversity (Kuuluvainen et al. 2002). tive management in terms of restoration is widely recog- Disturbance dynamics are also used as a guideline in devel- nized (Angelstam 1998; Kouki et al. 2001; Kuuluvainen et oping new forest management practices (Fries et al. 1997; al. 2002). Restoration, aiming to the rehabilitation of natural Bergeron et al. 2002; Franklin et al. 2002) that include, for structures, processes, and species composition in ecosystems example, the retention of living or dead trees and promoting altered by human actions (Bradshaw 1997), can be seen as a multilayered forest structure. tool to complement the traditional approach of conserving Saproxylic species, defined as being dependent on dead biodiversity by means of networks of reserves (Kouki et al. wood or on another dead wood dependent organism (Speight 2001; Kuuluvainen et al. 2002). Current restoration activity 1989), are a classic example of an ecological group that has in boreal forests is often based on imitating the disturbance been adversely affected by the extensive utilization of boreal

Received 18 September 2009. Accepted 19 December 2009. Published on the NRC Research Press Web site at cjfr.nrc.ca on 26 February 2010. T. Toivanen.1 Department of Biological and Environmental Science, University of Jyva¨skyla¨, P.O. Box 35, Jyva¨skyla¨ 40014, Finland. J.S. Kotiaho. Department of Biological and Environmental Science, University of Jyva¨skyla¨, P.O. Box 35, Jyva¨skyla¨ 40014, Finland; Natural History Museum, University of Jyva¨skyla¨, P.O. Box 35, Jyva¨skyla¨ 40014, Finland. 1Corresponding author (e-mail: [email protected]).

Can. J. For. Res. 40: 445–464 (2010) doi:10.1139/X09-205 Published by NRC Research Press 446 Can. J. For. Res. Vol. 40, 2010 forests (Grove 2002; Jonsson et al. 2005). The reasons that fecting the colonization and reproduction success of beetles: have led to the decline of these species include the reduction heavily charred wood hosts low densities (Saint- in the area of habitats (Axelsson and O¨ stlund 2001), the loss Germain et al. 2004) but even burnt logs (logs typically be- of connectivity between habitats (Komonen et al. 2000; Sii- come more charred than standing trees) may host high spe- tonen and Saaristo 2000), and the decline in the quality cies diversity and distinct species assemblages (Wikars (e.g., Fridman and Walheim 2000) and quantity (e.g., Siito- 2002; Gibb et al. 2006). An interesting aspect of burning is nen 2001) of dead wood. In particular, the absence of large- that it may reduce the resource specificity of saproxylic spe- diameter dead wood in managed forests and the changes in cies by enabling normally specialist species to utilize several disturbance dynamics (such as the prevention of forest fires tree species and thus making species assemblages of differ- and the removal of trees from storm damaged areas) can be ent tree species more similar (Wikars 2002). seen as major factors that have driven many saproxylic spe- Current restoration practices in boreal forests (Kuulu- cies to the edge of extinction. For example, in Finland, there vainen et al. 2002), of which controlled burning and creating are 183 threatened or extinct beetle species associated to dead wood by felling and damaging trees are most com- forests, of which 112 have been primarily affected by the monly used, are expected to improve habitat quality and in- loss of dead wood (Rassi et al. 2001). crease the availability of resources for saproxylic species. Several saproxylic species show resource specificity to a However, the effects of ongoing restoration actions are still particular tree genus, to a particular stage of wood decay, or inadequately understood. Here, we report results from a to a particular species of wood-decaying fungi (Jonsell et al. study in which we explored the relative importance of dif- 1998; Jonsson et al. 2005). For example, decaying deciduous ferent types of dead wood that had been created in restora- trees host assemblages that are distinct from those of coni- tion treatments for saproxylic beetles. Our main aims were fers. Within the boreal coniferous zone, the tree species to determine whether the abundance, species richness, and hosting most diverse assemblages is Norway spruce species assemblages of beetles differ between burnt and un- (Jonsson et al. 2005), but the role of aspen in hosting many burnt dead trees, how the increase in sun exposition due to rare or threatened saproxylic species is highlighted in sev- partial harvesting affects the beetle assemblages of the trees, eral studies (Jonsell et al.1998; Martikainen 2001; and how the responses of beetle assemblages to burning and Sverdrup-Thygeson and Ims 2002). The occurrence of the harvesting differ between two tree species. In addition, we majority of saproxylic invertebrates, and beetles in particu- studied whether down wood retention (DWR) had indirect lar, is often restricted to the first years following tree death effects on the beetle assemblages of the study trees via, for (Esseen et al. 1997; Jonsell et al. 1998). At the early decay example, fire intensity or affecting the attraction of beetles stages, the specialization of saproxylic invertebrates to a to the study sites. particular tree species is relatively high, but as the wood de- cay progresses, the tree species becomes less important. In Material and methods contrast, the role of wood-decaying fungi in determining the invertebrate community becomes more evident during the Study area later decay stages (Jonsell et al. 1998). The study was conducted on areas of a large-scale restora- Sun exposure is regarded as an important factor affecting tion experiment (Vanha-Majamaa et al. 2007) that were lo- species richness and species assemblages of saproxylic spe- cated in the vicinity of Evo, southern Finland (61811’N, cies. In natural conditions, sun-exposed habitats are formed 25805’E), within the south boreal vegetation zone. In total, by major disturbances such as forest fires and heavy storms. there were twenty-four 2 ha study plots located within a The majority of saproxylic beetles have been found to fa- 25 km  15 km area. The lands were owned by the Finnish vour sun-exposed substrates (e.g., Lindhe et al. 2005), and Forest and Park Service (six plots), Finnish Forest Research most of the red-listed saproxylic invertebrates in Sweden Institute (four plots), forest product company UPM (four have been classified to prefer sun-exposed sites or to be in- plots), Ha¨me Polytechnic University of Applied Sciences different to light conditions (Jonsell et al. 1998). In particu- (six plots), and the town of Ha¨meenlinna (four plots). All of lar, species associated with deciduous trees, e.g., birch and the plots were on average 80-year-old managed forest. The aspen, are likely to be adapted to sun-exposed conditions dominant tree species of the plots (~90% of the volume of (Kaila et al. 1997; Sverdrup-Thygeson and Ims 2002). How- standing trees) was Norway spruce (Picea abies (L.) Karst.) ever, recent evidence suggests that the importance of sun- with some birch (Betula spp.) and Scots pine (Pinus sylvest- exposed sites may have been overestimated to some extent ris L.). Before the experiment, the volume of standing trees because invertebrates tend to be more active in sun-exposed on the plots was 251.9 ± 64.8 m3/ha (mean ± SD) and the conditions, which may lead to sampling biases (Jonsell et al. volume of dead wood (diameter >5 cm for logs and >10 cm 2004). for stumps) was 17.3 ± 13.7 m3/ha; the volumes of living or Habitats created by forest fire are known to host high dead wood did not differ between the plots (Lilja et al. richness of saproxylic species and to be particularly impor- 2005). The dead wood at the plots consisted almost exclu- tant for rare and red-listed species (Simila¨ et al. 2002; Hy- sively of logging residues: small-diameter (<20 cm) logs va¨rinen et al. 2006; Toivanen and Kotiaho 2007a, 2007b). and cut stumps (Lilja et al. 2005). Relatively few species are directly dependent on burnt wood (Wikars 1997), but a wide range of saproxylic species Experimental design are attracted to the large quantities of dead wood that are Controlled burning and partial harvesting with DWR were available at sun-exposed conditions after forest fire (Siitonen applied as restoration treatments at the study plots to create 2001). Fire severity is likely to be an important factor af- habitats that mimic post-disturbance successional stages.

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During February and March 2002, 18 study plots were har- A) and formed four functional groups for the analyses. The vested such that the volume of standing trees was set to groups used were cambium consumers (cambium and 50 m3/ha and six plots were left unharvested. On the har- phloem feeders), associates of cambium consumers (mainly vested plots, 5, 30, or 60 m3/ha of cut down wood of predators but also species that are, for example, scavengers >10 cm diameter was retained (six plots of each treatment, in the burrows of bark beetles), fungivores, and wood borers referred to as harvest5, harvest30, and harvest60). The origi- (species that spend at least part of the larval development in nal relative abundances of the tree species were retained in the sapwood). We also formed a group of rare and red-listed the cuttings. saproxylic species. Rare species consisted of the beetle spe- In summer 2002, 12 plots (three plots of each harvesting cies that have been recorded in up to twenty-five 10 km  with DWR treatment and three unharvested plots) were 10 km squares in Finland (Rassi 1993) and red-listed species burnt. The first five burnings (two unharvested and three were the species considered threatened (IUCN categories harvest30 plots) were conducted in mid-June, the following CR, EN, and VU) or near threatened (NT) in Finland (Rassi five (three harvest5 and two harvest60 plots) in mid-July, et al. 2001). and the last two (one unharvested and one harvest60 plot) at the beginning of August. The fires were more intense at Statistical analyses the harvested plots causing high mortality of trees, while The response variables included were the species richness the mortality was quite low at the unharvested plots. Imme- and abundance of all saproxylic species and of the func- diately after the fire, the average volume of standing dead tional groups and the species richness of rare and red-listed trees was 1.4 m3/ha on unharvested plots, 24.1 m3/ha on har- saproxylic species. Moreover, we analysed the species-spe- vest5, 14.8 m3/ha on harvest30, and 43.8 m3/ha on harvest60 cific responses of the 50 most abundant species. This analy- (for detailed description of the treatments, see Lilja et al. sis was performed only if at least 30 individuals were 2005). collected, resulting in 49 species in year 2003 and 39 spe- We selected three even-sized Norway spruces (diameter at cies in 2004. The species abundance data were log10(x +1) breast height 23.1 ± 0.74 cm) and two birches (diameter transformed before the analyses. Each study tree was treated 21.5 ± 0.87 cm) as study trees at each study plot. At the as an individual observation, but because there were several burnt plots, we selected standing trees that had been killed trees of both species within one study plot, we analysed the by the fire. To enable the comparison of burnt and unburnt data with nested ANOVA. Burning, harvesting with DWR, dead trees of the same age, the selected standing trees on and tree species were entered into the model as fixed fac- unburnt plots were killed by girdling (stripping off a tors. Study plot was entered as random factor, nested within ~20 cm piece of bark around a tree at 1.3 m height to pre- burning  harvesting with DWR, and crossed with tree spe- vent water and nutrient flow) in the beginning of June 2002. cies. We analysed the data of the first and second post-treat- ment years separately. One girdled spruce had fallen during Sampling and grouping of beetles summer 2003 and it therefore was removed from the analy- During the 2 years following the treatments, we sampled ses. We performed the analyses with SPSS 15.0 for Win- beetles from the study trees with window traps that were at- dows software (SPSS Inc., Chicago, Illinois). tached to the tree trunks. The traps consisted of two cross- We used detrended correspondence analysis (DCA) with wise-set transparent plastic panes with a funnel and Canoco 4.0 (ter Braak 1987) to explore the compositional container below them. The upper edge of the funnel was variation in beetle assemblages. The species that occurred pressed against the tree trunk such that the length it touched in one sample only were excluded and the species abun- the trunk was about 25 cm. We used saline water with deter- dance data were log10(x + 1) transformed before analyses. gent in the containers to preserve the beetles. Here, we re- Detrending was performed using second-order polynomials. port data on beetles that were collected between 10 To statistically verify the clustering in the ordination, we an- May and 10 July 2003 (referred to as the first post-treatment alysed the loadings on axes 1 and 2 with nested ANOVA. year) and between 10 May and 10 July 2004 (referred to as the second post-treatment year). The activity of the majority Results of saproxylic beetle species peaks during early summer, and therefore, a representative sample of the beetle assemblages Abundance of saproxylic beetles can be collected within the given time period. In the first post-treatment year, we recorded 123 402 obli- We identified the majority of the beetles caught (99.98%) gatorily saproxylic beetles (93.8% of the trapped individu- to species. The identification of females of genera Philhygra als) representing 220 species (Appendix A). More (Staphylinidae) and Euplectus (Staphylinidae) was left to the individuals were collected on spruces than on birches; the genus level as a rule. In addition, we were not able to reli- abundance of cambium consumers and their associates was ably identify a few beetles of the genera Acrotrichis, Corti- higher on spruces but the abundance of fungivores and caria, , and Oxypoda. The nomenclature wood borers did not differ (Table 1). Burning did not affect follows Silfverberg (2004). the number of beetle individuals in general, but the abun- We extracted obligatorily saproxylic species (grouping dances of associates of cambium consumers, fungivores, based on literature such as Saalas (1917, 1923), Palm and wood borers were higher on burnt trees (Table 1). There (1951, 1959), and Koch (1989–1992), expert opinions, and was an interaction between burning and tree species our own experience) from the data and included only these (Table 1): less individuals were collected on burnt spruces species in the analyses. Furthermore, we classified the sap- than on unburnt spruces but more individuals on burnt roxylic species according to their larval ecology (Appendix birches than on unburnt birches (Fig. 1). This pattern was

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Table 1. ANOVA statistics for the effects of burning, harvesting with DWR and tree species on the abundance of all saproxylic beetles and of functional groups in the two post- treatment years.

Associates of cambium All saproxylic Cambium consumers consumers Fungal feeders Wood borers Variable Year df Fp df Fp df Fp df Fp df Fp Tree species 1 1,16 447.6 <0.001 1,16 628.4 <0.001 1,16 215.5 <0.001 1,16 0.278 0.605 1,16 0.972 0.339 2 1,16 245.6 <0.001 1,16 443.0 <0.001 1,16 64.78 <0.001 1,16 1.727 0.207 1,16 20.28 <0.001

Burning 1 1,16 3.054 0.100 1,16 0.434 0.519 1,16 30.12 0.001 1,16 17.01 0.001 1,16 12.83 0.002 2 1,16 56.98 <0.001 1,16 35.57 <0.001 1,16 77.70 <0.001 1,16 36.22 <0.001 1,16 18.55 0.001

Harvesting with DWR 1 3,16 7.035 0.003 3,16 6.317 0.005 3,16 4.721 0.015 3,16 1.645 0.219 3,16 6.114 0.006 2 3,16 2.052 0.147 3,16 1.704 0.206 3,16 2.349 0.111 3,16 1.821 0.184 3,16 10.21 0.001

Tree species  burning 1 1,16 61.81 <0.001 1,16 93.40 <0.001 1,16 24.57 <0.001 1,16 4.042 0.061 1,16 4.538 0.049 2 1,16 11.78 0.003 1,16 32.70 <0.001 1,16 0.417 0.528 1,16 0.030 0.865 1,16 1.763 0.203

Tree species  harvest- 1 3,16 2.575 0.090 3,16 3.343 0.046 3,16 6.867 0.003 3,16 3.277 0.048 3,16 0.540 0.661 ing with DWR 2 3,16 2.067 0.145 3,16 4.224 0.022 3,16 0.644 0.598 3,16 1.389 0.282 3,16 0.394 0.759

Burning  harvesting 1 3,16 0.194 0.899 3,16 1.074 0.388 3,16 0.180 0.908 3,16 0.989 0.423 3,16 1.812 0.186 with DWR 2 3,16 4.588 0.017 3,16 2.487 0.098 3,16 7.759 0.002 3,16 0.834 0.495 3,16 2.891 0.068

Tree species  1 3,16 1.159 0.356 3,16 1.766 0.194 3,16 3.322 0.047 3,16 1.061 0.393 3,16 0.275 0.842 burning  harvesting 2 3,16 4.013 0.026 3,16 7.207 0.003 3,16 2.599 0.088 3,16 1.581 0.233 3,16 3.113 0.056 with DWR

Plot (burning  1 16,13.3 2.647 0.040 16,12.6 2.596 0.047 16,13.4 2.897 0.028 16,13.8 5.948 0.001 16,15.0 3.516 0.010

harvesting with DWR) 2 16,14.8 0.763 0.701 16,14.3 1.756 0.146 16,14.4 0.466 0.929 16,14.3 3.116 0.019 16,14.1 2.240 0.068 2010 40, Vol. Res. For. J. Can. ulse yNCRsac Press Research NRC by Published Plot  tree species 1 16,71 0.825 0.654 16,71 0.657 0.826 16,71 0.860 0.615 16,71 1.022 0.445 16,71 2.300 0.009 2 16,71 1.845 0.041 16,71 1.368 0.183 16,71 1.689 0.069 16,71 1.173 0.311 16,71 1.223 0.273 Toivanen and Kotiaho 449

Fig. 1. Effect of burning, harvesting with DWR, and tree species on the number of saproxylic beetle individuals (mean ± SE) collected on individual trees in the two post-treatment years.

due to fire negatively affecting the abundances of cambium ers did not differ between burnt and unburnt spruces. Har- consumers and their associates on spruce, while the abun- vesting with DWR had no effect on the total number of dances of fungivores and wood borers were consistently individuals and the only functional group that was affected higher on burnt trees (Table 1). The total number of individ- was wood borers that were more abundant on trees at har- uals (Fig. 1) and the abundances of cambium consumers, vested plots than on those at unharvested plots (Table 1). In their associates, and wood borers were also affected by har- addition, there were interactions between burning and har- vesting with DWR (Table 1). This effect was due to that vesting with DWR and between burning, harvesting with more beetles were collected on trees at harvested plots than DWR, and tree species (Table 1). These interactions arose on those at unharvested plots (Tukey’s pairwise comparisons from harvesting having a negative effect on total number of for total number of individuals: unharvested versus harvest5, individuals (Fig. 1) and on the abundance of cambium con- p < 0.001; unharvested versus harvest30, p < 0.001; unhar- sumers and their associates among unburnt spruces. vested versus harvest60, p = 0.022). In addition, the total number of beetles collected was higher on trees at harvest5 Species richness of saproxylic beetles than on those at harvest60 (p = 0.004). The abundance of In the first year, the total number of saproxylic beetle spe- fungivores was not affected by harvesting with DWR cies (Fig. 2) and the species richness of cambium consum- (Table 1). ers, their associates, and wood borers were higher on In the second year, we recorded 40 795 obligatorily sap- spruce, while the species richness of fungivores tended to roxylic beetles (93.6% of the trapped individuals) represent- be higher on birch (Table 2). Total number of species ing 210 species (Appendix A). The total number of (Fig. 2) and the species richness of all functional groups saproxylic beetle individuals (Fig. 1) and the abundances of were higher on burnt trees (Table 2). There were interac- cambium consumers, their associates, and wood borers were tions between burning and tree species (Table 2): the differ- higher on spruce but the abundance of fungivores did not ence between burnt and unburnt trees was larger on birch differ between tree species (Table 1). The total number of (Fig. 2). Total number of species and the species richness individuals (Fig. 1) and the abundance of all functional of cambium consumers, their associates, and wood borers groups were higher on burnt trees but there were interactions were affected by harvesting with DWR (Table 2): more spe- between burning and tree species (Table 1): total number of cies were collected on trees at harvested plots than on those individuals (Fig. 1) and the abundance of cambium consum- at unharvested plots (Tukey’s pairwise comparisons for total

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Fig. 2. Effect of burning, harvesting with DWR, and tree species on the number of saproxylic beetle species (mean ± SE) collected on individual trees in the two post-treatment years.

number of species: p < 0.001 in all comparisons) (Fig. 2) ces than on birches but harvesting with DWR had no effect but the volume of DWR had no effect on the number of spe- (Table 2; Fig. 3). There was an interaction between burning cies collected. and tree species (Table 2): the number of rare and red-listed In the second year, the total number of saproxylic beetle species did not differ between burnt and unburnt spruces but species (Fig. 2) and the species richness of cambium con- more species were collected on burnt birches than on un- sumers were higher on spruce but the species richness of burnt ones. In addition, burnt spruces and birches did not wood borers did not differ between tree species and the spe- differ from each other but more rare and red-listed species cies richness of fungivores was higher on birch (Table 2). were collected on unburnt spruces than on unburnt birches Total number of species (Fig. 2) and the species richness of (Fig. 3). all functional groups were higher on burnt than on unburnt In the second year, we recorded 22 rare and red-listed trees (Table 2). The effect of burning was stronger on birch saproxylic species (including five NT species). Burning in- (interactions between burning and tree species, Table 1) creased the number of species but harvesting with DWR (Fig. 2). Harvesting with DWR affected the species richness and tree species had no effect and there were no interactions of wood borers (more species were collected on trees at har- between the factors (Table 2; Fig. 3). vested plots) but only tended to affect the total number of species and did not affect the species richness of other func- Composition of beetle assemblages tional groups (Table 2). However, there were interactions The DCA ordinations revealed significant differences in between burning and harvesting with DWR and between the composition of beetle assemblages. In the first post- burning, harvesting with DWR, and tree species (Table 2), treatment year (Fig. 4), there was distinct grouping accord- which were due to harvesting having a negative effect ing to tree species (on axis 1, F[1,15.99] = 298.9, p < 0.001; among unburnt spruces (Fig. 2). on axis 2, F[1,15.99] = 27.55, p < 0.001) and burning (on axis 1, F[1,15.98] = 27.48, p < 0.001; on axis 2, F[1,16.01] = 87.82, Species richness of rare and red-listed saproxylic species p < 0.001), and also harvesting with DWR affected the as- In the first post-treatment year, we recorded 26 rare and semblages (on axis 1, F[1,15.98] = 0.164, p = 0.919; on axis red-listed saproxylic species (including three VU and seven 2, F[1,16.01] = 9.292, p = 0.001). The effect of harvesting NT species). In general, the number of these species was in- with DWR was due to the assemblages of trees at harvested creased by burning and more species were collected on spru- plots differing from those at unharvested plots, but the vol-

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Table 2. ANOVA statistics for the effects of burning, harvesting with DWR, and tree species on the species richness of all saproxylic beetles, of functional groups, and of rare and red-listed saproxylic species in the two post-treatment years.

Associates of cambium All saproxylic Cambium consumers consumers Fungal feeders Wood borers Rare and red-listed Variable Year df Fpdf Fpdf Fpdf Fpdf Fpdf Fp Tree species 1 1,16 50.60 <0.001 1,16 90.87 <0.001 1,16 41.237 <0.001 1,16 4.400 0.052 1,16 9.884 0.006 1,16 12.50 0.003 2 1,16 7.932 0.012 1,16 40.85 <0.001 1,16 3.297 0.088 1,16 5.158 0.037 1,16 0.422 0.525 1,16 2.924 0.107

Burning 1 1,16 30.91 <0.001 1,16 17.41 0.001 1,16 29.76 <0.001 1,16 10.57 0.005 1,16 46.35 <0.001 1,16 7.683 0.014 2 1,16 64.38 <0.001 1,16 22.05 <0.001 1,16 34.38 <0.001 1,16 20.53 <0.001 1,16 40.72 <0.001 1,16 13.73 0.002

Harvesting with 1 3,16 4.236 0.022 3,16 4.063 0.025 3,16 4.056 0.025 3,16 0.975 0.429 3,16 8.794 0.001 3,16 0.668 0.584 DWR 2 3,16 3.216 0.051 3,16 0.220 0.881 3,16 0.894 0.466 3,16 2.380 0.108 3,16 15.88 <0.001 3,16 1.226 0.333

Tree species  1 1,16 8.001 0.012 1,16 6.552 0.021 1,16 16.76 <0.001 1,16 1.295 0.272 1,16 10.38 0.005 1,16 12.46 0.003 burning 2 1,16 5.307 0.035 1,16 7.048 0.017 1,16 5.865 0.028 1,16 0.096 0.760 1,16 0.069 0.796 1,16 0.602 0.449

Tree species  1 3,16 0.186 0.904 3,16 1.544 0.242 3,16 1.128 0.368 3,16 1.414 0.275 3,16 0.690 0.571 3,16 2.791 0.074 harvesting with 2 3,16 0.918 0.455 3,16 1.045 0.400 3,16 0.358 0.784 3,16 1.075 0.388 3,16 0.439 0.728 3,16 1.935 0.165 DWR

Burning  1 3,16 0.192 0.900 3,16 0.653 0.593 3,16 0.335 0.800 3,16 0.623 0.610 3,16 4.483 0.018 3,16 1.427 0.272 harvesting with 2 3,16 4.016 0.026 3,16 0.770 0.528 3,16 4.548 0.017 3,16 1.964 0.160 3,16 3.706 0.034 3,16 1.221 0.334 DWR

Tree species  1 3,16 0.896 0.465 3,16 0.992 0.421 3,16 0.851 0.486 3,16 0.848 0.488 3,16 0.415 0.744 3,16 1.106 0.376 burning  2 3,16 3.002 0.061 3,16 5.339 0.010 3,16 1.356 0.292 3,16 1.193 0.344 3,16 1.730 0.201 3,16 2.969 0.063 harvesting with DWR

Plot(burning  1 16,13.8 4.036 0.006 16,13.8 2.275 0.066 16,12.1 4.079 0.009 16,13.9 3.584 0.011 16,13.7 2.230 0.071 16,12.7 3.118 0.023 harvesting with 2 16,14.8 1.507 0.218 16,14.8 2.443 0.056 16,15.2 0.815 0.656 16,14.1 2.566 0.042 16,14,3 0.576 0.856 16,12.9 2.202 0.079 DWR) ulse yNCRsac Press Research NRC by Published

Plot  tree species 1 16,71 1.012 0.455 16,71 1.016 0.451 16,71 0.565 0.900 16,71 1.090 0.381 16,71 1.001 0.466 16,71 0.668 0.815 2 16,71 1.990 0.026 16,71 0.738 0.747 16,71 2.831 0.001 16,71 1.172 0.312 16,71 1.311 0.215 16,71 0.720 0.765 451 452 Can. J. For. Res. Vol. 40, 2010

Fig. 3. Effect of burning, harvesting with DWR, and tree species on the number of rare and red-listed saproxylic beetle species (mean ± SE) collected on individual trees in the two post-treatment years.

Fig. 4. DCA ordinations for the effects of burning, harvesting, and tree species on the composition of beetle assemblages in the two post- treatment years.

ume of DWR had no effect on the assemblages. Burning and suggesting that the assemblages of burnt birches and spruces tree species had a strong interaction (on axis 1, F[1,15.99] = were more similar to each other than those of unburnt 11.76, p = 0.003; on axis 2, F[1,15.99] = 12.54, p = 0.003), birches and spruces.

Published by NRC Research Press Toivanen and Kotiaho 453

Table 3. Effects of burning, harvesting, and tree species on the abundance of the 50 commonest beetle species collected in the two post- treatment years.

Abundance Abundance Burning Burning Harvest Harvest Tree Tree Additional Additional Species 2003 2004 2003 2004 2003 2004 2003 2004 2003 2004 Hylastes cunicularius Erichson 27 610 14 925 – 0 0 0 s s () Pityogenes chalcographus (Lin- 25 222 1 454 – 0 + 0 s s a naeus) (Curculionidae) Rhizophagus ferrugineus (Pay- 17 059 3 754 + + + 0 s s kull) (Monotomidae) Epuraea muehli Reitter (Nitidu- 11 058 32 – 0 0 0 s s b, c lidae) Dryocoetes autographus (Ratze- 6 353 5 012 + + + 0 s s a, b burg) (Curculionidae) Trypodendron lineatum (Olivier) 3 828 26 0 na 0 na s na (Curculionidae) Epuraea pygmaea (Gyllenhal) 3 375 315 + + 0 0 s 0 c (Nitidulidae) Hylastes brunneus Erichson 3 343 34 – 0 + 0 s s (Curculionidae) Crypturgus subcribrosus Eggers 2 922 6 605 – 0 0 0 s s b (Curculionidae) Hylastes opacus Erichson (Cur- 2 368 17 + na + na s na a culionidae) Ips typographus (Linnaeus) 2 006 445 – 0 + 0 s s b (Curculionidae) Pityophagus ferrugineus (Lin- 1 514 814 + + + 0 s s naeus) (Nitidulidae) Epuraea marseuli Reitter (Niti- 1 353 50 + 0 0 0 s 0 dulidae) Orthotomicus suturalis (Gyl- 1 142 94 + + 0 0 s s lenhal) (Curculionidae) Polygraphus poligraphus (Lin- 1 126 689 0 0 0 – s s naeus) (Curculionidae) Hylurgops palliatus (Gyllenhal) 1 049 18 0 na 0 na s na (Curculionidae) Ampedus nigrinus (Herbst) (Ela- 874 444 + + + + 0 0 teridae) Pityophthorus micrographus 561 29 – na 0 na s na b (Linnaeus) (Curculionidae) Crypturgus cinereus (Herbst) 552 1 103 0 + 0 0 s s b a (Curculionidae) Glischrochilus quadripunctatus 506 76 + 0 + 0 0 0 (Linnaeus) (Nitidulidae) binotata Reitter (Nitidu- 503 712 + + + 0 s s lidae) Orthoperus rogeri Kraatz (Cory- 498 58 + + 0 0 s 0 b b lophidae) Hylobius abietis (Linnaeus) 471 75 + + 0 0 s 0 b (Curculionidae) Thanasimus formicarius (Lin- 444 22 + na + na s na naeus) (Cleridae) Ips amitinus (Eichhoff) (Curcu- 430 19 – na + na s na b, c lionidae) Xylita laevigata (Hellenius) 415 513 + + + + b s a () Enicmus rugosus (Herbst) (Latri- 381 136 – 0 0 + 0 0 b diidae) Melanotus castanipes (Paykull) 363 332 + 0 + 0 0 0 (Elateridae) Trypodendron signatum (Fabri- 353 82 + 0 0 0 b b cius) (Curculionidae) Xylechinus pilosus (Ratzeburg) 344 27 – na – na s na (Curculionidae) Hylecoetus dermestoides (Lin- 339 115 + + 0 0 b b naeus) (Lymexylidae) Rhagium inquisitor (Linnaeus) 323 98 + 0 + 0 s 0 c (Cerambycidae)

Published by NRC Research Press 454 Can. J. For. Res. Vol. 40, 2010

Table 3 (concluded).

Abundance Abundance Burning Burning Harvest Harvest Tree Tree Additional Additional Species 2003 2004 2003 2004 2003 2004 2003 2004 2003 2004 Orthotomicus laricis (Fabricius) 260 15 + na 0 na s na (Curculionidae) Epuraea rufomarginata (Ste- 239 49 + 0 0 0 b b phens) (Nitidulidae) Quedius xanthopus Erichson 202 89 – – – – b 0 (Staphylinidae) Phloeonomus sjo¨bergi Strand 179 117 + 0 0 0 s s a (Staphylinidae) Hylobius pinastri (Gyllenhal) 167 39 0 0 + 0 b 0 (Curculionidae) Agathidium seminulum (Lin- 152 180 – – 0 0 0 0 naeus) (Leiodidae) Placusa atrata (Mannerheim) 140 7 + na 0 na 0 na c (Staphylinidae) Phloeostiba lapponica (Zetter- 139 2 + na 0 na 0 na stedt) (Staphylinidae) Nudobius lentus (Gravenhorst) 136 73 + + + + 0 0 (Staphylinidae) Plegaderus vulneratus (Panzer) 119 95 0 0 + 0 s s (Histeridae) Placusa incompleta Sjo¨berg 114 3 – na 0 na s na (Staphylinidae) Rhagium mordax (de Geer) 111 80 0 – + 0 b b (Cerambycidae) Xylita livida (Sahlberg) (Melan- 102 31 0 0 0 0 s s dryidae) Litargus connexus (Fourcroy) 99 52 + + 0 0 b b (Mycetophagidae) Salpingus ruficollis (Linnaeus) 98300000bb (Salpingidae) Ampedus balteatus (Linnaeus) 74 107 + + + 0 0 0 (Elateridae) Dacne bipustulata (Thunberg) 54 107 + + + + b b (Erotylidae) Sphaeriestes stockmanni (Bis- 17 90 na + na + na 0 tro¨m) (Salpingidae) Note: The ANOVA statistics are not shown but the symbols refer to significant (p < 0.05) effects. Burning and harvesting: +, positive effect; 0, no effect; –, negative effect. Tree species: 0, no preference; s, spruce; b, birch. Additional effect: a, negative response to the volume of DWR; b, interaction between burning and harvesting; c, interaction between burning and tree species. na, not analysed.

In the second post-treatment year (Fig. 4), the assemb- lionidae)), species living inside dead wood (e.g., Melanotus lages were more heterogeneous but there was still distinct castanipes (Paykull) (Elateridae) and Xylita laevigata (Hel- grouping according to tree species (on axis 1, F[1,16.01] = lenius) (Melandryidae)), or fungal feeders (e.g., Agathidium 111.2, p < 0.001; on axis 2, F[1,16.02] = 0.792, p = 0.387) seminulum (Linnaeus) (Leiodidae)) remained stable or even and burning (on axis 1, F[1,16.01] = 16.78, p = 0.001; on axis increased in the second post-treatment year (Table 3). 2, F[1,16.01] = 39.50, p < 0.001) and a weaker effect accord- The preferences of individual species were analysed only ing to harvesting with DWR (on axis 1, F[1,16.01] = 1.810, if at least 30 individuals were collected. In the first post- p = 0.186 ; on axis 2, F[1,16.01] = 4.269, p = 0.021). The in- treatment year, 27 of the 49 analysed species favoured burnt teraction of burning and tree species was still strong (on axis trees, the abundance of nine species did not differ between 1, F[1,16.01] = 23.45, p < 0.001; on axis 2, F[1,16.02] = 0.205, burnt and unburnt trees, and 13 species favoured unburnt p = 0.657). trees. The effect of harvesting with DWR on species’ prefer- ences was due to responses to harvesting in most cases, Species-specific responses while the actual volume of DWR had only minor effects. The abundance of the majority of the analysed species de- Twenty-one species were more abundant on trees at har- creased between the first and the second post-treatment vested plots, 26 species were indifferent to harvesting, and years, the decrease being most obvious among primary colo- only two favoured trees on unharvested plots. Four species nizers that are phloem feeders of recently died or dying trees had a negative response to the volume of DWR, but none (e.g., Ips spp. De Geer (Curculionidae) and Pityogenes chal- responded positively. In addition, nine species showed an in- cographus (Linnaeus) (Curculionidae)) and their associates teraction between burning and harvesting such that they fav- (e.g., Epuraea muehli Reitter (Nitidulidae) and Placusa spp. oured burnt trees at unharvested plots but unburnt trees at Erichson (Staphylinidae)). In contrast, the abundance of sec- harvested plots. Thirty species favoured spruce and 10 spe- ondary bark beetles (e.g., Crypturgus spp. Erichson (Curcu- cies birch, while the abundance of nine species did not differ

Published by NRC Research Press Toivanen and Kotiaho 455 between the tree species. Four species showed an interaction Effect of burning and tree species on the abundance of between burning and tree species such that they favoured beetles spruce among unburnt trees but were indifferent or favoured The effect of burning on the abundance of saproxylic spe- birch among burnt trees (Table 3). cies was dependent on the tree species and changed with In the second year, 17 of the 39 analysed species favoured time. In the first post-treatment year, more beetles were col- burnt trees, the abundance of 19 species did not differ be- lected on unburnt spruces than on burnt ones. This was tween burnt and unburnt trees, and only three species fav- mostly due to the responses of cambium consumers, primary oured unburnt trees. Seven species favoured trees on bark beetles in particular, that dominated the assemblages of harvested plots, while 30 species were indifferent to harvest- recently killed spruces. For example, the numbers of P. ing and two species favoured trees on unharvested plots. chalcographus and Ips typographus (Linnaeus) (Curculio- Two species had a negative response to the volume of nidae) were negatively affected by fire. These species have DWR. Only two species showed a similar interaction be- been found to avoid fire-scorched logs, most likely because tween burning and harvesting as in the first year. Seventeen the logs desiccate very rapidly and burning reduces the nu- species favoured spruce and seven species birch, while the tritional quality of the cambium (Wikars 2002; Gibb et al. abundance of 15 species did not differ between the tree spe- 2006; Johansson et al. 2006, 2007). The deteriorating effect cies. One species showed an interaction between burning of fire is not so strong on standing trees because only the and tree species (Table 2). base of the tree gets heavily charred, but it is nevertheless likely that the standing burnt spruces do not provide as Discussion good resource for the phloem-feeding primary colonizers as the unburnt spruces. However, the majority of the beetle General pattern of succession and methodological notes species were more abundant on burnt than on unburnt spru- This study covers the succession of beetle assemblages ces, and following the collapse of the numbers of primary during the first 2 years following tree death. Since disturb- colonizers, the total abundance of beetles was also higher ance-favouring species are likely to have good dispersal on burnt spruces in the second year. abilities (Southwood 1962; Jonsson et al. 2005) and to colo- In contrast with spruces, more beetles were consistently nize disturbance areas very rapidly (Hyva¨rinen et al. 2005), collected on burnt birches than on unburnt ones. The nega- the colonization of primary bark beetles and their associates tive effect of fire on the quality of cambium may not be so probably started immediately after the treatments. Therefore, strong in birch because the thicker bark makes birch more beetles collected during the first post-treatment year are fire resistant than spruce. In addition, the number of likely to have consisted of the offspring of the first coloniz- cambium consumers and their associates in birch is low ers and of the secondary species that had just started to ar- compared with that in spruce but the beetle assemblages of rive. In the second post-treatment year, the abundance of birch are to a larger extent determined by the fungal com- many of the primary species had already decreased strongly, munity. While the abundances of other functional groups while the abundance of, for example, species that develop were higher on spruce than on birch, the abundance of fun- inside dead wood or fungal feeders remained stable or even givores was not affected by tree species. Burnt trees are rap- increased. The decline of the primary colonizers can be ex- idly colonized by ascomycete fungi such as moulds (Penttila¨ plained by the fast depletion of the phloem layer of the and Kotiranta 1996; Wikars 2002) and by some ‘‘ruderal’’ trees, but the abundance of beetles may also have been af- species of corticoid fungi that can take advantage of the fected by adverse weather conditions. Compared with competition-free substrate (Penttila¨ and Kotiranta 1996). summer 2003, summer 2004 was colder and rainier, which Thus, the burnt birches are likely to provide a larger variety may have led to lower activity of beetles and affected the of resources for saproxylic species than unburnt birches dur- reproductive success. Hence, one must be careful in compar- ing the first years after tree death. ing the absolute numbers between the study years. Samples collected with window traps attached to standing Effect of burning and tree species on species richness and trees are not accurate measures of reproductive success of species assemblages beetles in a particular tree, but the traps also measure the The total species richness of saproxylic beetles and that of colonization and attraction of beetles to the trees. In addi- all functional groups were consistently higher on burnt trees tion, the traps may accidentally collect beetles that are not than on unburnt trees, but the effect was stronger among associated with the particular dead wood resource. Hence, birches than among spruces. Since burning decreased the the results reflect the beetle assemblages of the whole sur- abundance of the most common primary colonizers, the rounding area to some extent and they are likely to slightly higher species richness may have been brought about by re- underestimate the differences between different types of duced competition. Burnt trees may also provide more vari- dead wood (also see Wikars et al. 2005). However, the ma- able resources, or a wider range of microhabitats, for jority of beetles collected were obligatorily saproxylic spe- saproxylic beetles. For example, many saproxylic beetles cies that are also likely to reproduce in the particular trees. are known to depend on ascomycete fungi (Crowson 1984) There were distinct differences between tree species within that increase after fire, and these species also include some the study plots that have the same general beetle assemblage true fire specialists (Wikars 1997). On the other hand, our and the results are in concordance with the known ecology experiment does not enable distinguishing the effect of burnt of the individual saproxylic species. Therefore, we conclude substrate from the effect of burnt environment that may be that the results provide indirect evidence of the quality of more favourable for the reproduction of saproxylic beetles individual trees as a resource for saproxylic beetles. and to which more beetles may be attracted. With regard to

Published by NRC Research Press 456 Can. J. For. Res. Vol. 40, 2010 restoration in practice, however, there is no need to separate the abundance of beetles and only tended to affect total spe- these two effects because as a rule, burnt wood is available cies richness. The species richness of wood borers was still at a burnt environment only. higher on trees at harvested plots, while the species richness The numbers of rare and red-listed species were relatively and abundance of other groups on unburnt spruces were low on the study trees and most of the species (except for E. negatively affected by harvesting. Therefore, it appears that muehli, which was very abundant on spruces during the first although unburnt spruces at harvested areas attract high year) occurred on one or two study trees only. The effect of numbers of beetles immediately after tree death, the trees burning was consistently positive among birches, while desiccate extremely fast, which leads to a rapid decline of more rare and red-listed species were collected on burnt species dependent on the phloem layer, i.e., cambium con- spruces than on unburnt ones in the second post-treatment sumers and their associates. At unharvested areas, spruces year only. Among burnt trees, there was no difference be- may retain their moisture content for a longer time and pro- tween tree species, but unburnt spruces attracted more rare vide a longer-lasting resource for primary species. In addi- and red-listed species than unburnt birches in the first year. tion, many spruce-living species are likely to be adapted to This suggests again that burning is particularly effective in shaded conditions because spruce forests are characterized creating habitats for birch-living species but the spruce-liv- by small gap dynamics and long fire rotation rather than fre- ing species may not benefit from burning that much. quent large-scale disturbances (Esseen et al. 1997; Kuuluvai- The beetle assemblages were distinct between burnt and nen 2002; Wikars 2002). However, it must be noted that the unburnt trees and, as expected, between spruces and birches. unburnt spruces were killed by girdling, which may cause Interestingly, the assemblages collected on burnt spruces and abnormally fast tree death, and thus, the succession of beetle birches were more similar to each other than those collected assemblages in girdled trees may differ from that in natu- on unburnt spruces and birches. Normally monophagous rally formed dead wood. species may become less tree species specific when living in burnt wood (Wikars 2002), and there were indeed some Effect of DWR species that were collected on burnt trees of the ‘‘wrong’’ The volume of dead wood may affect species richness via species in our study. However, significant changes in the providing a wider array of habitats, allowing the buildup of preferences of beetle species were rare. Thus, the similarity larger populations with lower risk of extinction or providing of beetle assemblages between burnt birches and spruces temporal continuity of suitable habitats (Siitonen 2001). was more likely due to the beetle assemblages of burnt trees However, the amount of felled trees of the same size and being to a larger extent dominated by generalist species than decay stage is unlikely to have direct effects on the beetle those of unburnt trees (Wikars 2002). Species colonizing re- assemblages of individual trees within a short time period. cently dead trees usually show strong tree species specific- As expected, the effect of the volume of DWR was gener- ity, while the importance of tree species decreases as decay ally weak, but in the first post-treatment year, it was nega- progresses (Jonsell et al. 1998; Jonsson et al. 2005), but spe- tively associated with the number of beetles collected. At cies favouring burnt trees may be more associated with a unburnt plots, felled trees provide an obvious resource for particular species of fungus or with the burnt environment saproxylic beetles, and therefore, fewer beetles may have itself than with a particular tree species. been attracted to individual standing trees when there were numerous host trees available. The negative effect of DWR Effect of harvesting and the dependence of the effect on was particularly evident among spruces, which is logical be- burning and tree species cause most of the felled trees were spruces. At burnt plots, Harvesting increased the abundance and species richness the value of the felled trees as a resource was probably of beetles on the study trees in the first post-treatment year. lower because most beetles are likely to avoid heavily This can be attributed to increased sun exposure that is charred logs. However, the logs provide burning fuel, and likely to be beneficial to several disturbance adapted beetle therefore, fire intensity increased with the volume of DWR. species. In particular, several primary colonizers are known This may have caused the negative effect of DWR on the to favour sun-exposed areas (Jonsell et al. 1998). The colo- number of beetles at burnt plots. nization of trees at harvested areas may also occur more rap- idly, which can be due to stronger attraction to the sites or Efficiency of restoration efforts that trees die more rapidly at open areas because of, for ex- Forest restoration that imitates natural disturbances is ample, increased drought. However, fungivores were not af- likely to be efficient only if the disturbance is typical of the fected by harvesting and it is likely that shaded conditions particular forest type and the target species are adapted to are more favourable for fungal growth. In addition, some the particular disturbance. In spruce-dominated forests, primary species were positively affected by harvesting at un- large-scale disturbances such as forest fires are likely to be burnt plots but negatively affected at burnt plots. This pat- naturally scarce and unpredictable (Kuuluvainen 2002), and tern is likely due to fire intensity being low at unharvested hence, a substantial part of the species inhabiting dead spru- plots compared with harvested plots. The reproductive suc- ces may not be adapted to open post-disturbance habitats. cess of has been found to decrease with increasing This may be particularly true for late successional species fire severity, although there is much variation depending on with weak dispersal abilities (Southwood 1962). Neverthe- the tree species (Saint-Germain et al. 2004). Norway spruce less, burning appears to be an efficient way to increase spe- has very thin bark, and thus, strong effects of fire intensity cies richness of saproxylic beetles within a short time period can be expected. (also see Hyva¨rinen et al. 2006; Toivanen and Kotiaho In the second year, harvesting had no general effect on 2007b). However, while birch-living species consistently

Published by NRC Research Press Toivanen and Kotiaho 457 benefit from burning, the effect of burning is weak, and in Fries, C., Johansson, O., Pettersson, B., and Simonsson, P. 1997. some cases negative, among spruce-living species. Trees at Silvicultural models to maintain and restore natural stand struc- harvested areas attract high numbers of beetles immediately tures in Swedish boreal forests. For. Ecol. Manag. 94(1–3): 89– after the disturbance, but the resource may prove short- 103. doi:10.1016/S0378-1127(97)00003-0. lasting. Therefore, long-term studies are needed to clarify Gibb, H., Pettersson, R.B., Hja¨lte´n, J., Hilszczan´ski, J., Ball, J.P., the value of restoration efforts, and in particular to deter- Johansson, T., Atlegrim, O., and Danell, K. 2006. Conservation- mine whether late successional species are able to colonize oriented forestry and early successional saproxylic beetles: re- the disturbance areas. However, it should be noted that the sponses of functional groups to manipulated dead wood sub- aim of restoration is not only to provide resources for sap- strates. Biol. Conserv. 129(4): 437–450. doi:10.1016/j.biocon. roxylic species but also to facilitate the succession towards 2005.11.010. Grove, S.J. 2002. Saproxylic ecology and the sustainable more natural, multilayered forest structure. management of forests. Annu. Rev. Ecol. Syst. 33(1): 1–23. Acknowledgements doi:10.1146/annurev.ecolsys.33.010802.150507. Hyva¨rinen, E., Kouki, J., Martikainen, P., and Lappalainen, H. The restoration experiment was established by the Forest 2005. Short-term effects of controlled burning and green-tree re- Ecology section of the University of Helsinki, and we thank tention on beetle (Coleoptera) assemblages in managed boreal especially Timo Kuuluvainen, Saara Lilja, and Pasi Puttonen forests. For. Ecol. Manag. 212(1–3): 315–332. doi:10.1016/j. for their co-operation. The treatments were carried out by foreco.2005.03.029. the land owners and were helped by numerous students and Hyva¨rinen, E., Kouki, J., and Martikainen, P. 2006. Fire and volunteers. Jarno Nevalainen and Satu Kuntsi carried out a green-tree retention in conservation of red-listed and rare dead- substantial part of the beetle identification. Merja Aho, Veli wood-dependent beetles in Finnish boreal forests. Conserv. Liikanen, and Jarno Nevalainen helped in the field work and Biol. 20(6): 1710–1719. doi:10.1111/j.1523-1739.2006.00511.x. the material was sorted by Satu Kuntsi and Elina Manninen. PMID:17181806. Juha Siitonen provided valuable information for the classifi- Johansson, T., Gibb, H., Hilszczanski, J., Pettersson, R.B., Hja¨lte´n, cation of beetles. This research was funded by the MOSSE J., Atlegrim, O., Ball, J.P., and Danell, K. 2006. Conservation- research program of the Ministry of Agriculture and For- oriented manipulations of coarse woody debris affect its value estry, Otto A. Malm’s Foundation, Jenny and Antti Wihuri’s as habitat for spruce-infesting bark and ambrosia beetles (Co- leoptera: Scolytinae) in northern Sweden. Can. J. For. Res. Foundation, Koneen Sa¨a¨tio¨ Foundation, Maj and Tor Nes- 36(1): 174–185. doi:10.1139/X05-235. sling’s Foundation, and The Finnish Centre of Excellence in Johansson, T., Hja¨lten, J., Gibb, H., Hilszczanski, J., Stenlid, J., Evolutionary Research. Ball, J.P., Alinvi, O., and Danell, K. 2007. Variable response of different functional gropus of saproxylic beetles to substarte ma- References nipulation and forest management: implications for conservation Angelstam, P.K. 1998. Maintaining and restoring biodiversity in strategies. For. Ecol. Manag. 242(2–3): 496–510. doi:10.1016/j. European boreal forests by developing natural disturbance re- foreco.2007.01.062. gimes. J. Veg. 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Table A1. Saproxylic species collected in the two post-treatment years, their total abundance, and the number of study sites (n = 24) where they were recorded.

First post-treatment year Second post-treatment year Species Ecology Status Abundance Sites Abundance Sites Carabidae Dromius agilis (Fabricius) P, gen 15 12 6 5 Dromius quadraticollis Morawitz P, gen Rare 2 1 Histeridae Plegaderus saucius Erichson P, con 1 1 Plegaderus vulneratus (Panzer) P, gen 119 19 95 20 Platysoma angustatum (Hoffmann) P, gen 28 6 3 2 Platysoma deplanatum (Gyllenhal) P, gen 23 7 Platysoma lineare Erichson P, con 18 6 Platysoma minus (Rossi) P, gen 6 3 Ptiliidae Pteryx suturalis (Heer) F, gen 1 1 Leiodidae Anisotoma axillaris Gyllenhal F, gen 2 2 1 1 Anisotoma castanea (Herbst) F, gen 1 1 1 1 Anisotoma glabra (Kugelann) F, gen 26 15 36 13 Anisotoma humeralis (Fabricius) F, gen 8 7 20 13 Agathidium confusum Brisout de Barneville F, gen 5 4 14 7 Agathidium nigripenne (Fabricius) F, gen 15 11 16 8 Agathidium rotundatum (Gyllenhal) F, gen 2 2 4 3 Agathidium seminulum (Linnaeus) F, gen 152 21 180 24 Scydmaenidae Stenichnus bicolor (Denny) P, gen 3 3 1 1 Microscydmus nanus (Schaum) P, gen 1 1 Staphylinidae Xylostiba monilicornis (Gyllenhal) P, gen 3 3 Phloeostiba lapponica (Zetterstedt) P, gen 139 17 2 1 Phloeostiba plana (Paykull) P, gen 6 4 Phloeonomus pusillus (Gravenhorst) P, gen 14 6 Phloeonomus sjobergi Strand P, gen 179 22 117 23 Bibloporus bicolor (Denny) P, gen 5 4 11 10 Bibloporus minutus Raffray P, gen Rare 2 1 Euplectus kirbii Denny P, gen Rare 1 1 Euplectus mutator Fauvel P, gen Rare 1 1 Euplectus piceus Motschulsky P, gen 5 3 3 3 Euplectus punctatus Mulsant P, gen 2 2 4 4 Tyrus mucronatus (Panzer) P, gen 2 2 4 3 Phloeocharis subtilissima Mannerheim P, gen 2 2 Sepedophilus constans (Fowler) F, con 1 1 7 5 Sepedophilus littoreus (Linnaeus) F, gen 96 22 14 11 Sepedophilus testaceus (Fabricius) F, gen 38 12 12 5 Pentanota meuseli Bernhauer P, dec Rare 4 2 Phloeopora corticalis (Gravenhorst) P, gen 21 11 16 10 Phloeopora testacea (Mannerheim) P, gen 66 21 14 10 Dadobia immersa (Erichson) P, gen 6 6 4 4 Dinaraea aequata (Erichson) P, gen 13 7 4 3 Gyrophaena boleti (Linnaeus) F, gen 5 4 4 3 Leptusa pulchella (Mannerheim) P, gen 1 1 2 2 Euryusa castanoptera Kraatz P, dec 15 9 3 3 Anomognathus cuspidatus (Erichson) P, gen 18 10 17 9 plana (Gyllenhal) P, gen 27 11 3 3 Placusa atrata (Mannerheim) P, gen 140 20 7 4 Placusa complanata Erichson P, con 25 7 2 2

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Table A1 (continued).

First post-treatment year Second post-treatment year Species Ecology Status Abundance Sites Abundance Sites Placusa depressa Ma¨klin P, con 51 16 1 1 Placusa incompleta Sjo¨berg P, dec 114 20 3 3 Placusa suecica Johnson & Lundberg P, con Rare 1 1 Placusa tachyporoides (Waltl) P, gen 61 15 3 3 Scaphisoma agaricinum (Linnaeus) F, gen 26 15 44 14 Scaphisoma assimile Erichson F, dec 1 1 Scaphisoma subalpinum Reitter F, gen 1 1 1 1 Nudobius lentus (Gravenhorst) P, con 136 21 73 17 Atrecus affinis (Paykull) P, con 6 5 Gabrius expectatus Smetana P, con 14 9 9 4 Bisnius subuliformis (Gravenhorst) P, gen NT 3 2 Quedius maurus (Sahlberg) P, gen 4 4 Quedius plagiatus Mannerheim P, gen 6 4 10 7 Quedius xanthopus Erichson P, gen 202 18 89 14 Lucanidae caprea (DeGeer) F/W, dec 48 16 30 14 Buprestidae Melanophila acuminata (DeGeer) W, gen, pyro NT 25 7 3 3 Anthaxia quadripunctata (Linnaeus) W, con 7 3 Elateridae Lacon conspersus (Gyllenhal) P/W, gen NT 1 1 3 2 Denticollis linearis (Linnaeus) P/W, gen 1 1 Ampedus balteatus (Linnaeus) P/W, gen 74 20 107 18 Ampedus erythrogonus (Mu¨ller) P/W, gen 37 12 63 16 Ampedus nigrinus (Herbst) P/W, gen 874 24 444 23 Ampedus pomonae (Stephens) P/W, gen 1 1 Ampedus suecicus Palm P/W, gen NT 2 2 Ampedus tristis (Linnaeus) P/W, gen 10 6 21 12 Melanotus castanipes (Paykull) P/W, gen 363 24 332 23 Lycidae Dictyoptera aurora (Herbst) P, gen 14 10 9 8 Lygistopterus sanguineus (Linnaeus) P, gen 1 1 Cantharidae Malthodes brevicollis (Paykull) P/W, gen 1 1 Malthodes pumilus (Brebisson) P/W, gen 1 1 Dermestidae Globicornis emarginata (Gyllenhal) D, gen 2 2 Megatoma undata (Linnaeus) D, gen 1 1 3 3 Anobidae Ptinus subpillosus Sturm F/D, gen 46 17 3 2 abietis (Fabricius) C, con 13 9 5 3 Ernobius explanatus (Mannerheim) C, con 9 7 1 1 Ernobius mollis (Linnaeus) C, con 1 1 5 2 pertinax (Linnaeus) W, gen 3 2 18 10 Dorcatoma punctulata Mulsant & Rey F, gen 1 1 Dorcatoma robusta Strand F, dec 1 1 2 2 Lymexylidae Hylecoetus dermestoides (Linnaeus) C, gen 339 24 115 18 Hylecoetus flabellicornis (Schneider) C, con 72 5 14 5 Trogossitidae Peltis grossa (Linnaeus) F, gen NT 1 1 4 4 Ostoma ferruginea (Linnaeus) F, gen 4 3 3 3 Nemozoma elongatum (Linnaeus) P, con Rare 4 3

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Table A1 (continued).

First post-treatment year Second post-treatment year Species Ecology Status Abundance Sites Abundance Sites Cleridae Thanasimus femoralis (Zetterstedt) F, gen 10 8 10 5 Thanasimus formicarius (Linnaeus) F, gen 444 22 22 9 Aplocnemus impressus (Marsham) P, gen Rare 1 1 Aplocnemus tarsalis (Sahlberg) P, con 6 5 Dasytes niger (Linnaeus) P, gen 10 7 49 10 Sphindidae Sphindus dubius (Gyllenhal) F, gen 4 4 6 6 Aspidiphorus orbiculatus (Gyllenhal) F, gen 12 7 10 7 Nitidulidae Epuraea angustula Sturm P/F, gen 85 19 52 10 Epuraea biguttata (Thunberg) F, gen 15 7 8 4 Epuraea boreella (Zetterstedt) P/F, gen 6 6 5 5 Epuraea deubeli Reitter P/F, con Rare 1 1 Epuraea laeviuscula (Gyllenhal) P/F, con 19 8 Epuraea marseuli Reitter D/F, gen 1 353 23 50 14 Epuraea muehli Reitter P/F, con Rare 11 058 24 32 12 Epuraea neglecta (Heer) D/F, dec Rare 2 1 2 1 Epuraea oblonga (Herbst) P/F, con 45 14 16 7 Epuraea pallescens (Stephens) D/F, gen 6 4 Epuraea pygmaea (Gyllenhal) D/F, gen 3 375 24 315 16 Epuraea rufomarginata (Stephens) D/F, gen 239 22 49 16 Epuraea thoracica (Tournier) D/F, con 2 1 Epuraea unicolor Olivier F, gen 2 1 1 1 Epuraea variegata (Herbst) F, gen 3 2 Reitter F, gen 503 22 712 24 Glischrochilus hortensis (Geoffroy) F, dec 12 7 5 4 Glischrochilus quadripunctatus (Linnaeus) F, gen 506 22 76 10 Pityophagus ferrugineus (Linnaeus) P, con 1 514 24 814 24 Monotomidae Rhizophagus bipustulatus (Fabricius) P, gen 8 7 2 2 Rhizophagus cribratus Gyllenhal P, dec 3 1 1 1 Rhizophagus depressus (Fabricius) P, dec 17 10 1 1 Rhizophagus dispar (Paykull) P, gen 16 10 42 13 Rhizophagus ferrugineus (Paykull) P, gen 17 059 24 3 754 24 Rhizophagus nitidulus (Fabricius) P, gen 3 3 6 4 Rhizophagus parvulus (Paykull) P, gen 46 18 24 8 Silvanidae Dendrophagus crenatus (Paykull) P, gen 17 10 10 8 Silvanoprus fagi (Guerin-Me´ne´ville) P, con 21 11 6 5 Laemophloidae Laemophloeus muticus (Fabricius) F, dec, pyro 1 1 1 1 Cryptolestes abietis (Wankowicz) P, con 29 9 79 17 Cryptolestes alternans (Erichson) P, con 23 7 52 14 Henoticus serratus (Gyllenhal) F, gen, pyro 42 9 Pteryngium crenatum (Fabricius) F, gen 1 1 Micrambe longitarsis (J.Sahlberg) F, con Rare 10 21 4 3 Cryptophagus corticinus Thomson F, dec, pyro Rare 2 5 2 2 Atomaria bella Reitter F, gen 14 9 30 14 Atomaria subangulata J.Sahlberg F, gen 1 1 6 6 Atomaria umbrina (Gyllenhal) F, gen 3 3

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Table A1 (continued).

First post-treatment year Second post-treatment year Species Ecology Status Abundance Sites Abundance Sites Erotylidae Dacne bipustulata (Thunberg) F, dec 54 13 107 14 Tritoma bipustulata Fabricius F, dec Rare 2 2 2 2 Triplax aenea (Schaller) F, dec 1 1 Triplax russica (Linnaeus) F, dec 6 6 8 6 fagi Brisout de Barneville F, dec 2 2 3 2 Cerylon ferrugineum Stephens F, gen 3 3 2 2 Cerylon histeroides (Fabricius) F, gen 13 9 22 10 Endomychidae Endomychus coccineus (Linnaeus) F, dec 3 1 Corylophidae Clypastraea pusilla (Gyllenhal) F/D, gen, pyro VU 1 1 Orthoperus rogeri Kraatz F/D, gen 498 24 58 21 Latridiidae Latridius hirtus Gyllenhal F, gen 9 8 5 4 Enicmus fungicola Thomson F, gen 23 13 11 4 Enicmus planipennis Strand F, gen Rare 50 16 2 2 Enicmus rugosus (Herbst) F, gen 381 23 136 23 Corticaria crenicollis Mannerheim F, gen Rare 11 4 1 1 Corticaria lapponica (Zetterstedt) F, con 1 1 Corticaria lateritia Mannerheim F, dec 7 6 9 7 Corticaria polypori J.Sahlberg F, con Rare 1 1 1 1 Mycetophagidae Litargus connexus (Geoffroy) F, gen 99 16 55 16 Mycetophagus fulvicollis Fabricius F, gen Rare 1 1 Mycetophagus piceus (Fabricius) F, gen 2 2 16 10 Ciidae Cis boleti (Scopoli) F, dec 8 5 17 8 Cis glabratus Mellie´ F, gen 3 2 1 1 Cis hispidus (Paykull) F, dec 47 17 24 16 Cis micans (Fabricius) F, dec NT 4 3 Cis punctulatus Gyllenhal F, gen 11 6 4 3 Orthocis alni (Gyllenhal) F, gen 12 10 15 11 Hadreule elongatula (Gyllenhal) F, gen Rare 1 1 4 3 Sulcacis affinis (Gyllenhal) F, dec 1 1 9 8 Melandryidae Orchesia fasciata (Illiger) F, gen 1 1 1 1 Orchesia micans (Panzer) F, gen 1 1 Orchesia minor Walker F, gen 1 1 Xylita laevigata (Hellenius) F/W, gen 415 23 513 24 Xylita livida (Sahlberg) F/W, gen 102 20 31 16 Zilora ferruginea (Paykull) F, con 1 1 aculeata Linnaeus F, dec 1 1 Zopheridae Bitoma crenata (Fabricius) P, gen 5 3 22 9 Tenebrionidae Bius thoracicus (Fabricius) F, gen Rare 1 1 Corticeus linearis (Fabricius) P, con 33 14 11 8 Corticeus suturalis (Paykull) P, con Rare 1 1 3 3 Bolitophagus reticulatus (Linnaeus) F, dec 37 10

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Table A1 (continued).

First post-treatment year Second post-treatment year Species Ecology Status Abundance Sites Abundance Sites Oedemeridae Calopus serraticornis (Linnaeus) C, gen 26 12 29 15 Chrysanthia geniculata Heyden C, con 1 1 Pythidae Pytho depressus (Linnaeus) C, con 72 15 1 1 Pyrochroidae Schizotus pectinicornis (Linnaeus) F, gen 2 2 1 Salpingidae Rabocerus foveolatus (Ljungh) P, dec 1 1 1 1 Rabocerus gabrieli (Gerhardt) P, dec 4 4 7 5 Sphaeriestes bimaculatus (Gyllenhal) P, con 1 1 Sphaeriestes stockmanni (Bistro¨m) P, gen, pyro NT 7 7 90 9 Salpingus ruficollis (Linnaeus) P, gen 98 18 30 16 Aderidae Euglenes pygmaeus (DeGeer) F, dec 2 1 Anidorus nigrinus (Germar) F, dec Rare 1 1 12 7 Scraptiidae Anaspis bohemica Schilsky P, dec 2 2 Anaspis marginicollis Lindberg P, gen 12 8 Cerambycidae Asemum striatum (Linnaeus) W, gen 39 13 7 5 Tetropium castaneum (Linnaeus) C, con 86 16 17 7 Tetropium fuscum (Fabricius) C, con 1 1 Rhagium inquisitor (Linnaeus) C, gen 323 21 98 21 Rhagium mordax (DeGeer) C, gen 111 21 80 22 Oxymirus cursor (Linnaeus) W, gen 5 5 4 3 Pachyta lamed (Linnaeus) C, con 8 6 virginea (Linnaeus) C, con 1 1 2 2 pratensis (Laicharting) C/W, con 10 5 38 10 Alosterna tabacicolor (DeGeer) W, gen 1 1 Judolia sexmaculata (Linnaeus) C, gen 2 2 Molorchus minor (Linnaeus) C, gen 23 10 11 4 Callidium violaceum (Linnaeus) C, con 1 1 Xylotrechus rusticus (Linnaeus) C, dec NT 2 2 Pogonocherus fasciculatus (DeGeer) C, con 35 13 11 8 Acanthocinus aedilis (Linnaeus) C, con 2 2 Tropideres dorsalis (Thunberg) F, dec, pyro VU 1 1 Platyrhinus resinosus (Scopoli) F, dec, pyro VU 1 1 (Linnaeus) F, dec 6 4 15 7 Curculionidae Rhyncolus ater (Linnaeus) W, gen 4 3 4 3 Magdalis carbonaria (Linnaeus) C, dec 1 1 Magdalis violacea (Linnaeus) C, gen 1 1 Hylobius abietis (Linnaeus) C, gen 471 24 75 21 Hylobius piceus (DeGeer) C, con 15 10 5 4 Hylobius pinastri (Gyllenhal) C, con 167 24 39 16 Pissodes castaneus (DeGeer) C, con 1 1 Pissodes gyllenhali (Sahlberg) C, con 40 12 9 5 Pissodes harcyniae (Herbst) C, con 20 6 4 4 Pissodes pini (Linnaeus) C, con 15 8 7 4 hispidus (Linnaeus) C, dec 1 1 2 1 Hylurgops glabratus (Zetterstedt) C, con 3 3

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Table A1 (concluded).

First post-treatment year Second post-treatment year Species Ecology Status Abundance Sites Abundance Sites Hylurgops palliatus (Gyllenhal) C, con 1 049 24 18 11 Hylastes brunneus Erichson C, con 3 343 24 34 14 Hylastes cunicularius Erichson C, con 27 610 24 14 925 24 Hylastes opacus Erichson C, con 2 368 22 17 11 Xylechinus pilosus (Ratzeburg) C, con 344 12 27 6 Tomicus piniperda (Linnaeus) C, con 7 5 Dendroctonus micans (Kugelann) C, con 1 1 1 1 Phloeotribus spinulosus (Rey) C, con 9 5 6 6 Polygraphus poligraphus (Linnaeus) C, con 1 122 17 689 19 Polygraphus subopacus Thomson C, con 4 3 Scolytus ratzeburgi Janson C, dec 4 1 5 3 Pityogenes bidentatus (Herbst) C, con 15 9 3 3 Pityogenes chalcographus (Linnaeus) C, con 25 222 24 1 545 24 Pityogenes quadridens (Hartig) C, con 14 7 2 1 Orthotomicus laricis (Fabricius) C, con 260 19 15 7 Orthotomicus proximus (Eichhoff) C, con 1 1 Orthotomicus suturalis (Gyllenhal) C, con 1 142 19 94 12 Ips amitinus (Eichhoff) C, con 430 22 19 15 Ips typographus (Linnaeus) C, con 2 006 23 445 23 Dryocoetes alni (Georg) C, dec 1 1 Dryocoetes autographus (Ratzeburg) C, con 6 353 24 5 012 24 Crypturgus cinereus (Herbst) C, con 552 21 1 103 22 Crypturgus hispidulus Thomson C, con 1 1 60 8 Crypturgus pusillus (Gyllenhal) C, con 22 10 60 16 Crypturgus subcribrosus Eggers C, con 2 922 24 6 505 24 Trypodendron domesticum (Linnaeus) C, dec 1 1 1 1 Trypodendron lineatum (Olivier) C, con 3 828 24 26 14 Trypodendron signatum (Fabricius) C, dec 353 22 82 13 Xyleborus dispar (Fabricius) C, dec 27 14 1 1 Cryphalus saltuarius Weise C, con 12 8 3 3 Pityophthorus lichtensteinii (Ratzeburg) C, con 6 2 Pityophthorus micrographus (Linnaeus) C, con 561 20 29 14 Note: Ecology: C, cambium consumer; D, detritivore; F, fungivore; P, predator; W, wood borer; gen, generalist (no strict preference for tree species); con, species of coniferous trees; dec, species of deciduous trees; pyro, fire-dependent species. Status: NT, near threatened; VU, vulnerable; rare, rare species in Finland.

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