Insect Conservation and Diversity (2011) doi: 10.1111/j.1752-4598.2011.00149.x Habitat requirements of the endangered beetle Boros schneideri (Panzer, 1796) (Coleoptera: Boridae) _ EREŠK IE N E_ and VIDMANTAS KARALIUS LAIMA BLAŽ YT E-Č Institute of Ecology of Nature Research Centre, Vilnius, Lithuania Abstract. 1. To further knowledge regarding habitat requirements of the threatened species Boros schneideri, a total of 1522 dead standing Scots pine (Pinus sylvestris) were checked for the presence of B. schneideri larvae. For the 245 trees inhabited by larvae (16% of all examined trees), tree characteristics and occupancy patterns were measured. 2. As most of the investigated forests were 40–80 years old and were dominated by comparatively thin dead trees (5–20 cm diameter at breast height), B. schneideri larvae were mainly found under the bark of trees with a diameter in the range of 10–20 cm. However, the probability of a tree being inhabited by B. schneideri increased progressively with tree diameter and forest age. 3. Most of the trees with B. schneideri (79%) were in medium-dense and sparsegrowth pine forests where the canopy cover was 60–80%. The presence of Norway spruce (Picea abies) in Scots pine forests was an important factor affecting the probability of trees being inhabited by B. schneideri. Only 6% of trees inhabited by B. schneideri were found in mixed Scots pine-Norway spruce forests, where the shadowing was higher than 80%. 4. All dead trees inhabited by B. schneideri had loose and at least slightly fragmented bark. A bark area of 0.08 m2 was found to be sufficient for the survival of B. schneideri. The critical bark thickness for B. schneideri larvae was 5 mm. 5. The data obtained are important for the optimisation of conservation measures implemented during forest management operations. Key words. Boridae, Boros schneideri, conservation, habitat, Pinus sylvestris, saproxylic beetle. Introduction Dead wood is one of the most important substrata for maintaining biodiversity in forest ecosystems (Esseen et al., 1997). Intensive forest management has reduced the volume of dead wood and its diversity in many regions (Kaila et al., 1997; Simila et al., 2003). As a result, due to a lack of suitable breeding substratum, many saproxylic species (species that are dependent on decaying wood) have declined, and in the last few decades many have become threatened (Jonsell et al., 1998; Tikkanen et al., 2006). Numerous ecological studies aimed at conserving the biodiversity of saproxylic species have been conducted (Ranius & Jansson, 2000; Jonsson et al., 2005; Lindhe et al., 2005; Johansson, 2006; Tikkanen et al., 2007; Alexander, 2008). Correspondence: Laima Blažyt_e-Čereškien_e, Institute of Ecology of Nature Research Centre, Akademijos 2, LT-08412 Vilnius21, Lithuania. E-mail: blazyte@ekoi.lt Success of the European network of conservation areas Natura 2000 depends on broad biological knowledge of the designated protected species. To devise management plans and develop protected areas, it is necessary to carry out in-depth analysis of habitat requirements of the target species. For this purpose, studies of habitat requirements of a few endangered saproxylic beetle species, such as Osmoderma eremita (Scopoli, 1763) (Cetoniidae) (Ranius & Nilsson, 1997), Cerambyx cerdo (Linnaeus, 1758) (Cerambycidae) (Buse et al., 2007), Pytho kolwensis (Sahlberg, 1833) (Pythidae) (Siitonen & Saaristo, 2000) and Cucujus haematodes (Erichson, 1845) (Cucujidae) (Horak et al., 2010) have been conducted. Boros schneideri (Panzer, 1796) is the only species in the family Boridae occurring in Europe (Levkaničovi˛, 2009). It is an endangered saproxylic beetle species listed in Annex 2 of the European Union Habitat Directive (Council Directive, 2006) and occurs mostly in the Baltic area. The species has decreased in number over the past century. B. schneideri is extremely rare in the  2011 The Authors Insect Conservation and Diversity  2011 The Royal Entomological Society 1 2 _ Laima Blazˇyte-Čeresˇkien e_ and Vidmantas Karalius Alpine biogeographical region (Lokality Natura, 2000). In the Boreal region, it is recorded in Scandinavia (Väisänen et al., 1993; Siitonen et al., 1999; Hyvärinen et al., 2005) and in the Baltic countries (Vilks & Telnov, 2003; Ferenca, 2003; Šablevičius, 2003; Sifverberg, 2004; Karalius et al., 2006; Karalius & Blažyt_e-Čereškien_e, 2009 and other). In the Continental region, the species has been reported only from Poland (Burakowski et al., 1987; Kubisz, 2004) and the Czech Republic (Chobot, 2005). In all these regions, the overall assessment of this species is unfavourable-inadequate or unfavourable-bad (Habitats Directive, 2009). A study of B. schneideri distribution in Lithuania revealed that this species is restricted to areas of forest greater than 20 000 ha (Karalius & Blažyt_e-Čereškien_e, 2009). It is likely that the current nature reserves of suitable habitat are too small for the long-term survival of this species, therefore adjacent unprotected forests should be managed in a sympathetic manner to create large blocks of suitable habitat. Saproxylic species may persist in managed forests only if a sufficient amount of dead wood is retained during forest management operations (Tikkanen et al., 2007). To manage the habitat for the survival of B. schneideri, it is necessary to know the sorts of wood suitable for this species as well as the most important substratum properties necessary for its successful reproduction. Unfortunately, data on habitat requirements and biology of B. schneideri are scarce and contradicting (Kubisz, 2004; Tikkanen et al., 2007). It is supposed that a generation may take at least 2 years to develop. The larval nutrition is still poorly studied. Some studies have indicated that they may be carnivorous and feed on other insect larvae (Kubisz, 2004). Other researchers have thought that larvae may feed on Aureobasidium mycelium (Uselis, 2008). The larvae pupate during July and August, with the adult beetles emerging in October and then overwintering under the bark of a tree, thereafter to be found until May of the following year (Uselis, 2008). It is known that dead Scots pine trees provide the main habitat for this species in mixed and boreal forests (Šablevičius, 2003; Karalius et al., 2006; Tikkanen et al., 2007; Karalius & Blažyt_e-Čereškien_e, 2009). The aims of this study were to examine and discuss easily measured characteristics of those Scots pine trees on which B. schneideri were found beneath the bark, thereby extending the knowledge regarding habitat requirements of this threatened species. Methods and study sites Study sites The study area (about 54–55N, 22–25E) is located at the boundary between the hemiboreal and temperate vegetation zones in Lithuania (Ahti et al., 1968). Investigations were carried out in forests dominated by Scots pine (Pinus sylvestris L.). Test plots (0.2–1 ha, depending on the biotope range) were selected in middle-aged (40- to 60-year-old) and older coniferous forests dominated by Scots pine (P. sylvestris L.), where Scots pines constituted the greatest proportion of the total volume of the growing stock, with smaller proportions of Norway spruce (Picea abies (L.) Karst.) and birch (Betula spp.). Ground-layer vegetation was dominated by Vaccinium myrtillus L., V. vitisidaea L., lichens, or moss. Such types of forest are known as B. schneideri dwelling sites (Karalius et al., 2006; Karalius & Blažyt_e-Čereškien_e, 2009). The fieldwork began in May 2006 when 200 dead trees in 13 localities in southern and eastern parts of Lithuania (Var_ena, Švenčionys, Taurag_e, and Jurbarkas districts) were examined. In June–October 2007, a further 943 dead trees were checked in 29 localities in southern, eastern and central parts of Lithuania (Taurag_e, Jurbarkas, Kaunas, Jonava, Lazdijai, Var_ena, Šalčininkai, Trakai, and Švenčionys districts), while in September 2008, an additional 379 dead trees in eight localities in the southern part of Lithuania (Var_ena district) were checked. Host trees and larvae sampling Dead standing P. sylvestris trees were selected based on previous studies (Karalius et al., 2006) which indicated that suitable trees for B. schneideri are greater than 5 cm in diameter at breast height (about 120 cm above the ground level), with loose and at least partially fragmented bark and a damp under-bark layer. All such trees in the plots were registered and checked for the presence of B. schneideri larvae by temporarily removing up to 1 ⁄ 2 of the bark from the test area (tree trunk surface at a height of up to 2 m above the ground level). Dead trees with the bark firmly stuck to the trunk, as well as trees with a dry under-bark layer were not included into the account. Inspection of a tree ceased when at least one B. schneideri specimen was detected. When such specimens were found, the temporarily opened bark was reattached to the trunk with small nails. Boros schneideri adults and larvae were identified by analysing their external morphology (Saalas, 1937; Bei-Bienko, 1965). Larvae are strongly flattened, subparallel, lightly sclerotized, yellowish and 17–23 mm in length (Levkaničovi˛, 2009). They can be identified as their abdomen is covered with chitin spikes that are crown-like and visibly different to larvae of other genera (Cucujus, Schizotus, Pytho) which are common under the bark of Scots pine. The body length and width of the larval head capsule was measured in field conditions. The measured larvae were released back onto the host tree. Only trees with B. schneideri larvae (and not with adults) were assumed to be inhabited by this species and were used for analysis. As the area of the opened bark varied in different cases, a potential larval density per square meter of bark was calculated. Bark area calculations were conducted assuming that the trunk was a 2 m high cylinder. A number of variables describing the characteristics of dead trees were recorded. The diameter of every checked tree was measured. For those trees inhabited by B. schneideri, additional characteristics recorded were tree height, tree canopy cover, percentage of the remaining bark on the trunk, percentage of the bark opened during inspection, the number of larvae found, larval position in relation to cardinal points and thickness of the bark at the larval spot on trees (Table 1). The modified Braun-Blanquet method was used to describe the degree of presence of Norway spruce in Scots pine forests (Balevičien_e, 1991). The cover of Norway spruce was  2011 The Authors Insect Conservation and Diversity  2011 The Royal Entomological Society, Insect Conservation and Diversity Habitat requirements of the endangered beetle Boros schneideri 3 Table 1. Variables measured from each of dead trees inhabited by Boros schneideri. Variable Range Explanation Diameter* Tree height 5–54 cm 10–35 m Canopy cover 5–90% Bark cover Opened bark Thickness of the bark No. of larvae found Larval position 10–95% 10–50% 4.7–35.7 mm Diameter at breast height (1.2 m above the ground level) Tree height was assessed approximately by actually measuring 1 ⁄ 10 of the tree height, which was determined from a distance with a measuring stick, and then multiplying it by 10 The sky hemisphere was mentally divided into northern and southern parts. The percentage of sky overshadowed by trees and bushes was evaluated for both parts Cover of the remaining bark on the tree trunk at a height of up to 2 m above the ground level Cover of the opened bark on the tree trunk at a height of up to 2 m above the ground level Thickness of the bark at the larval position was calculated as the mean of three independent measurements Number of larvae found under bark of one trunk Larval position in relation to cardinal points: N = northern, S = southern, W = western, E = eastern sides of the trunk 1–9 specimens N, S, W, E *The diameter of every checked tree was measured. determined visually, where: 0–+ = no or rare Norway spruce trees (1–2), 1–2 = cover 5–25%, 3 and more = cover more than 25%. Data relating to the age of stands was obtained from the local forestry office only in the investigations in 2007 and 2008. Statistical analysis Analysis was performed using the StatSoft, Inc. (Tulsa, OK, USA). The Shapiro–Wilk test was used for normality testing (Čekanavičius & Murauskas, 2000). The significance of the stand age and trunk diameter in explaining the probability of a trunk being inhabited by B. schneideri was studied by logistic regression. The Spearman correlation was used to estimate the relationship between different characteristics of a dead tree’s trunk and larval density. The Mann–Whitney U test was used to estimate differences in the percentage of trees inhabited by B. schneideri in Scots pine forests with different densities of Norway spruce. Results Occurrence of B. schneideri Larvae of B. schneideri were found under the bark of 245 dead trees from a sample of 1522 studied (16% of cases). The total number of larvae found was 335, the size of which varied from small (body length 4.0 mm, width of head capsule 0.54 mm) to large (body length 28.0 mm, width of head capsule 2.14 mm). The number of individuals recorded under bark varied from one to nine. The proportion of trees with one larva was the highest (71% of cases), as inspection of the tree ceased when at least one B. schneideri specimen was detected. The area of opened bark varied from 0.02 m2 to 1.13 m2 in different cases. Therefore, a potential larval density per square meter of bark at a height of up to 2 m above the ground level was calculated. The calculated larval density varied from 1.8 to 44.9 specimens per m2 (average 12.2  0.95 per m2). Larval density negatively correlated with the area of remaining bark (R = )0.54; P < 0.001) and with the area of opened bark (R = )0.78; P < 0.001) i.e. the density of larvae was higher on trees with a smaller area of bark cover. A bark area of 0.08 m2 was found to be sufficient for the survival of B. schneideri larva. The distribution of larvae in relation to cardinal points was similar: 24% of larvae were recorded on the southern side of the trunk, 24% on the eastern, 23% on the northern and 29% on the western. No significant correlations were found between larval density and any other parameters of dead trees. Host tree characteristics Boros schneideri larvae were recorded on many kinds of trunks according to the values of the trunk variables measured (Table 2). Trees with a diameter of 5–20 cm constituted most (80.3%) of the potential host trees. The abundance of trees thicker than 20 cm in diameter was inversely proportional to their diameter. The narrowest tree on which B. schneideri larvae were found under the bark had a diameter of 8 cm, while the thickest was 50 cm in diameter. Most trees with B. schneideri (54.1%) had a diameter of 10–20 cm. However, according to the logistic regression, the probability of a trunk being inhabited by B. schneideri increased with increasing diameter (Table 3). The height of trees with B. schneideri larvae varied from 10 m to 35 m. The average diameter of inhabited trunks was 18  0.5 m. The greatest proportion (42%) of trees was 15–20 m high. About 7% of dead trees inhabited by B. schneideri were broken snags with their height ranging from 4 to 10 m. Bark cover in the test area of trees with B. schneideri varied from 10 to 95%. Less than 10% bark coverage on the trunk was found to be insufficient for B. schneideri to survive. Bark thickness at the larval spot varied from 4.7 to 35.7 mm (average 15.1 mm) (Table 2). About 2% of larvae were found under bark thinner than 5 mm. Most larvae (77%) were found under bark 5–20 mm thick. Consequently, bark thickness of 5 mm was considered the minimum for the survival of B. schneideri larvae.  2011 The Authors Insect Conservation and Diversity  2011 The Royal Entomological Society, Insect Conservation and Diversity 4 _ Laima Blazˇyte-Čeresˇkien e_ and Vidmantas Karalius Table 2. Trunk variables in the sample dead trees: number of trunks from which the variable was measured, minimum, maximum, mean and median values. Variable N Minimum Maximum Mean Median Diameter (cm)* Tree height (m) Canopy cover (%) Bark cover (%) (m2) Opened bark (%) (m2) Thickness of the bark (mm) 245 (944) 165 165 159 8 (5.1) 4 5 10 0.08 10 0.02 4.7 50 (54) 35 90 95 2.84 50 1.13 35.7 19.9 (14.7) 18.3 65 56 0.74 32 0.19 15.1 17 (12.7) 18 70 60 0.67 30 0.14 14.5 159 159 *Figures in parenthesis are for the trunks without Boros schneideri. Table 3. A logistic regression model including trunk diameter and stand age which contributed significantly to the probability of a tree being inhabited by Boros schneideri. Variables R2 R P Regression equation Trunk diameter Stand age 0.8364 0.9146 0.0006 y = )797.462 + 8.012 · x 0.7184 0.8476 0.0039 y = 13.017 + 5.194 · x Canopy cover of trees being inhabited by B. schneideri varied from 5% in clear-cut plots to 90% in mixed pine-spruce forests (Table 2). Most of the studied trees with B. schneideri (79%) were in medium-dense and sparse-growth pine forests where the canopy cover varied from 60 to 80% (average 65  1.2%). Only 6% of trees inhabited by B. schneideri were found in mixed Scots pine-Norway spruce forests, where the shadowing was higher than 80%. The presence of Norway spruce in Scots pine forests was an important factor affecting probability of trees being inhabited by B. schneideri. The percentage of dead trees with B. schneideri larvae in Scots pine forests (Fig. 1) without or with few Norway spruce trees was statistically higher than in forests where Norway spruce trees covered more than 25% of the area (U = 80, Z = 2.2, P = 0.026). Thus, the probability of dead trees being inhabited by B. schneideri seems to be higher in Scots pine forests without shadowing by Norway spruce trees. The age of stands in the study areas varied from 40 to 127 years (average age 71  2.2 years). Most trees with B. schneideri (70%) were in 40- to 80-year-old forests. Only 30% of dead trees were in forests older than 80 years. However, the logistic regression showed that the probability of a dead tree being inhabited by B. schneideri increased with increasing stand age (Table 3). Discussion Boros schneideri is usually described as a species of relict primeval forests inhabiting old and thick dead trees (Telnov, 2003; Kubisz, 2004; European rarities in Estonia: Boros schneideri, 2007). Our data provide a substantial basis for the revision of Fig. 1. Probability of dead Scots pine trees being inhabited by Boros schneideri in Scots pine forests with different density of Norway spruce. The cover of Norway spruce was determined by Braun-Blanquet method, where 0–+ = no or few trees, 1–2 = 5–25% cover, 3 and more = more than 25% cover. Different letters (a, b) denote statistically significant differences (P < 0.05; Mann–Whitney U test). this view. It is true that B. schneideri prefers breeding on thick trees, which could be evidenced by the results of logistic regression. A greater B. schneideri population size on thicker trees might be attributed to the bark area, which is generally greater on thick trees than on thin trees. However, beetles can also inhabit relatively thin trees, which are 10–15 cm in diameter. As the abundance of dead trees with a diameter of 10–20 cm in managed forests was significantly higher than the abundance of thicker trees, thin dead trees in this 10–20 cm diameter range hosted the greatest proportion of B. schneideri population. In literature, it is indicated that tree size influences species richness of saproxylic beetles (Ranius & Jansson, 2000) and that trunk diameter can positively correlate with density of some saproxylic insect species and negatively with some other species (Lindhe et al., 2005). Our findings emphasize the importance of  2011 The Authors Insect Conservation and Diversity  2011 The Royal Entomological Society, Insect Conservation and Diversity Habitat requirements of the endangered beetle Boros schneideri comparatively thin trees as breeding substrata for B. schneideri. Consequently, dead trees with a diameter of 10–20 cm should be regarded as an important dead wood resource in B. schneideri conservation projects. According to Buse et al. (2007), bark thickness of trees is one of the significant predictors for the presence of saproxylic beetle. Our study shows that bark 7 mm thick provides ample conditions for the survival of B. schneideri larva in the forest. Optimal conditions for saproxylic beetle larvae would probably be large trunks with thick bark and phloem, these creating better conditions for larval development over a longer time period (Siitonen & Saaristo, 2000). However, the greatest proportion of the B. schneideri population in Lithuania seems to occur under middle thickness bark, because most large forests suitable for B. schneideri (Karalius & Blažyt_e-Čereškien_e, 2009) today consist of thin and medium diameter trees with a middle bark thickness. Boros schneideri larvae were found only under the bark of trees with loose bark, where the process of bark loss had already started. The density of B. schneideri larvae was higher under the bark of trees with smaller areas of remaining bark. This can be explained by the specificity of the bark loss process, which is usually accelerated by woodpeckers and in most cases proceeds from the top of the trunk downwards, because bark at the top is thinner and easier to remove. Larvae of B. schneideri develop under the bark for several years. As a result of bark loss, they are forced to congregate in smaller under-bark areas. Species richness and the number of red-listed saproxylic beetles are found to be significantly higher in sun-exposed trees (Lindhe & Lindelöw, 2004; Lindhe et al., 2005). The impact of canopy cover on a dead tree’s suitability for B. schneideri was found to be very important too. B. schneideri is defined as a type of species that lives in pine snags and favours warm and sunny stands (Tikkanen et al., 2007). Our data confirm that the species often inhabits 100% of dead biodiversity trees remaining in clear-cuts and maximally exposed to the sun. Alongside light preference, B. schneideri also demonstrates considerable tolerance to shady places: the species is able to breed under the bark of dead trees in medium-dense pine forests. Due to the scarcity of dead trees in open places, most of the population lives under the bark of dead trees in the shaded conditions of medium-dense and sparse-growth pine forests. However, dead trees are quite rarely inhabited by B. schneideri in dense Scots pine forests, especially with higher Norway spruce densities. The data obtained demonstrate which characteristics of dead wood have the highest value for B. schneideri conservation and are important for the optimization of conservation measures during forest management operations. To sum up, dead Scots pine trees with a diameter exceeding 10 cm, with loose and at least slightly fragmented bark, which stand in places with shadowing conditions varying from medium-dense pine forests with low Norway spruce densities to fully sun-exposed clear-cuts, are suitable breeding substrata for this species. Such trees should be retained during forest management operations, especially during thinning and cleaning. When planning conservation measures, special attention should be paid to the finding that, though B. schneideri beetles prefer breeding on thick and old trees, the main part of the population in Lithuania 5 occurs under the bark of dead trees with a diameter of 10–20 cm in comparatively young medium-dense and sparse-growth pine forests. Further research is needed to better understand if the probabilities of dead tree being inhabited by B. schneideri differ between forest types. Acknowledgement We thank Mr Vytautas Uselis and Ms Asta Uselien_e, ecologists at Viešvil_e Strict Nature Reserve, Mr Vincas Slavickas, a biologist at Dzukija National Park, Dr. Bronius Šablevičius, an ecologist at Aukštaitija National Park, and Mr Audrius Aliukonis and Mr Romas Ferenca, ecologists at Labanoras Regional Park for their kind of assistance during fieldwork. 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