ZooKeys 22: 309-340 (2009) A peer-reviewe: d open-access journal I doi: 10.3897/zookeys.22. 144 RESEARCH ARTICLE #Zookey S www.pens oftonline.n et/zoo keys Launched to accelerate biodiversity research Deadwood and saproxylic beetle diversity in naturally disturbed and managed spruce forests in Nova Scotia DeLancey J. Bishop'*, Christopher G. Majka’, Seren Bondrup-Nielsen?, Stewart B. Peck' | Department of Biology, Carleton University, Ottawa, Ontario, Canada 2 clo Nova Scotia Museum, 1747 Summer St., Halifax, Nova Scotia Canada 3 Department of Biology, Acadia University, Wolfville, Nova Scotia, Canada 4 RR 5, Canning, Nova Scotia, Canada Corresponding author: Christopher G. Majka (c.majka@ns.sympatico.ca) Academic editor: Jan Klimaszewski | Received 26 March 2009 | Accepted 6 April 2009 | Published 28 September 2009 Citation: Bishop DJ, Majka CG, Bondrup-Nielsen S, Peck SB (2009) Deadwood and saproxylic beetle diversity in naturally disturbed and managed spruce forests in Nova Scotia In: Majka CG, Klimaszewski J (Eds) Biodiversity, Bio- systematics, and Ecology of Canadian Coleoptera I. ZooKeys 22: 309-340. doi: 10.3897/zookeys.22.144 Abstract Even-age industrial forestry practices may alter communities of native species. Thus, identifying coarse patterns of species diversity in industrial forests and understanding how and why these patterns differ from those in naturally disturbed forests can play an essential role in attempts to modify forestry practices to minimize their impacts on native species. This study compares diversity patterns of deadwood habitat structure and saproxylic beetle species in spruce forests with natural disturbance histories (wind and fire) and human disturbance histories (clearcutting and clearcutting with thinning). We specifically examine how beetle diversity differs in relation to disturbance history and how beetle variation is linked to the di- versity of deadwood habitats. Beetle and deadwood data were collected from thirty spruce forests in Nova Scotia and analyzed under three related diversity perspectives: alpha (diversity within local forests); beta (heterogeneity among local forests within disturbance classes); and gamma (cumulative species richness within disturbance classes). Few data support a prediction of lower alpha deadwood and beetle diversity in managed forests, or a prediction of lower gamma species richness in managed forests. The beta scale analysis yielded support for the following two hypotheses: (1) beetle assemblages are different in forests with different disturbance histories; (2) turnover of beetle assemblages is higher among naturally disturbed forests than among managed forests. The prediction of lower gamma diversity of saproxylic beetle species in managed forests compared to naturally disturbed forests was not supported. The lack of differences between naturally disturbed and industrial forests in structures that are characteristic of older forests (e.g., large-diameter deadwood) may relate to the presence of residual deadwood in second growth forests lin- gering from before clearcut harvesting. However, such residual deadwood is only an artifact that will soon decay and not be replaced. This suggests that the continuity of deadwood microhabitats for species that depend on old-forest structures is only short-term. Copyright DeLancey J. Bishop et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 310 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Keywords Saproxylic, Coleoptera, deadwood, forest management, disturbance history, Nova Scotia, Canada Introduction Disturbance is one of the main determinants of ecosystem structure, composition, and function (Pickett and White 1985; Attiwill 1994; Abel and Stepp 2003). A major con- cern with respect to biodiversity loss in forest ecosystems is driven by the threat that disturbance imposed by modern forestry has on species diversity through altering the habitat structure (Haila and Kouki 1994; Haila et al. 1994; Siitonen 2001; Langor et al. 2006). Deadwood substrates serve as microhabitats that host an immense diversity of forest species (called saproxylics), primarily invertebrates (Speight 1989; Warren and Key 1991; Siitonen 1994, 2001, 2003; Grove 2002). Unfortunately, the physical di- versity of deadwood in forests has a high potential for being altered by modern forestry practices. Many saproxylic species are threatened due to loss of habitat (Siitonen 2001, 2003; Jonsson et al. 2005; Larsson et al. 2006, Langor et al. 2006). Thus, considering the fate of saproxylic species and their deadwood microhabitats in industrial forests is an essential part of any forest management initiative that values native species conservation and ecologically sustainable resource use (e.g., Canadian Council of Forest Ministers 1995; National Board of Forestry Sweden 1996; Canadian Forest Service 1997; Siito- nen 2001; Langor et al. 2006). The diversity of deadwood substrates (abundance and richness of physical states) is often changed when natural disturbances are replaced by forestry disturbances (Gore and Patterson 1986; Hagan and Grove 1996; Sturtevant et al. 1997; Siitonen 2001; Langor et al. 2006). Whereas natural disturbances occur stochastically, forestry is de- terministic (Attiwill 1994) and aimed toward specific economic ends, namely, the maximization of sustainable fiber productivity. Over recent decades, industry and gov- ernment in Canada and across the globe have favored even-age management practices of clearcut-replanting or clearcut-regeneration with thinning. Natural disturbances have therefore been largely replaced by even-age forestry. We predicted several changes to the local diversity of deadwood substrates in hu- man disturbed forests. Even-aged harvesting practices may drive (1) the loss of large diameter deadwood due to short time-spans of stand rotations; (2) a decreased local diversity of deadwood decay structures due to all deadwood being in a single decay co- hort which senesces through even-aged stand cycles; (3) the loss of standing deadwood due to the view that standing deadwood is a source of economic inefficiency and a safety hazard; and (4) the loss of deadwood from the many native tree species that are selected against because they are not considered economically valuable. We examined saproxylic beetles (Order Coleoptera) diversity patterns and rela- tionships with deadwood diversity patterns in managed and naturally disturbed spruce (Picea rubens Sarg. or Picea mariana Mill., Pinaceae) forests in Nova Scotia, Canada. Many species have highly specialized saproxylic niches (Speight 1989; Warren and Key Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 311 1991; Siitonen 2001; Langor et al. 2006), suggesting that species assemblages should be sensitive to variation in deadwood substrates. That most saproxylic beetles can be easily sampled using relatively inexpensive methods (WOkland 1996) favors the group logistically as well as ecologically. This study takes an exploratory look at a system that has only recently begun to receive detailed attention in North America (Hammond et al. 2001, 2004; Langor and Spence 2006; Langor et al. 2006). In the Maritime Provinces of Canada only Kehler et al. (2004) and Dollin et al. (2008) have previously examined saproxylic beetle com- munities in detail. Our basic objective is to identify whether modern forestry is gener- ating invertebrate diversity patterns that are different from those generated by natural disturbances. This task can be approached on many scales and from many perspectives on species diversity. Specifically, we adopt a three-part view on species diversity, similar to the alpha, beta, and gamma diversity perspectives proposed by Whittaker (1972). Alpha diversity pertains to local diversity or, in our case, diversity measured at the scale of a single forest stand. Beta diversity pertains to the turnover of alpha diversity between or among localities; in other words, the degree to which alpha diversity dif- fers from locality to locality. At the beta scale, we compare the turnover of species and deadwood diversity among local forests within four disturbance history classes (see methods). We use gamma diversity to refer to the total number of species cumulatively sampled within all forests in each disturbance history class. We conceptualized and tested the following set of specific predictions for the al- pha, beta, and gamma scales: Alpha scale: (1) naturally disturbed forests exhibit higher local diversity of deadwood (habitat) structures than forests that are under even-age management regimes; and (2) local diversity of saproxylic beetle assemblages within forests is proportional to the hetero- geneity of deadwood habitat structures within forests; and thus (3) naturally disturbed forests exhibit higher local diversity of saproxylic beetle species than forests that are under even-age management regimes. Beta scale: (1) beetle assemblages and forest structures are different in forests with different disturbance histories, which is to say that species and deadwood compositions are more similar within forest disturbance classes than among forest disturbance history class- es (see methods); (2) heterogeneity in deadwood habitat structures is higher among naturally disturbed forests than among forests that are under even-age management regimes; and (3) turnover of saproxylic beetle species among forests is proportional to the heterogeneity of deadwood habitat structures among forests; and thus (4) turnover of saproxylic beetle species is higher among naturally disturbed forests than among forests that are under even-age management regimes. Gamma scale: (1) the cumulative beetle species diversity across all naturally disturbed forests is higher than the cumulative beetle species diversity across all forests under even-age management regimes. 312 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Methods Study forests. Thirty spruce forests were selected for beetle sampling in central Nova Sco- tia, Canada, concentrated in two regions: the Liscomb Game Sanctuary (45° 09" N, 62° 30° W) and an area north of St. Margaret’s Bay (44° 44’ N, 63° 54’ W). The forests were chosen to represent one of four general disturbance histories, two anthropogenic and two natural: (1) clearcut origin (CC), (2) clearcut origin followed by pre-commercial or com- mercial thinning (CC+), (3) fire disturbance origin (F), and (4) wind disturbance origin (W/). We set several criteria for the experimental forests. Canopy trees had to be numeri- cally dominated (70 % minimum) by spruce (Picea spp.). Stands had to be large enough to contain a 40 x 120 m trap-sampling grid with a surrounding 100 m buffer (minimum of 7.7ha in total size) to limit edge effects from riparian areas, forests of different tree species composition, and any non-forested areas, including recent clearcuts. The closest two centers within forest were 800 m apart while the most distant two were separated by 190 km. Stands were in addition selected to reduce as much as possible geographic spatial dependence. Time since last intense disturbance (clearcut, fire, severe hurricane, etc.) was limited to a minimum of 30 years, as indicated by mean age of dominant canopy trees. Of the selected forests, mean stand ages ranged from 30 to over 250 years. A concen- tration of thinned forests in the young age range reflects the recent implementation of thinning as a management practice. Forest management in Nova Scotia was not strictly regulated and private woodlots constituting 75% of the province could be managed as the owners saw fit when the present study stands experienced their last disturbance. Thus, rotation ages vary greatly and the selection of stands could not be controlled with regard to ages of stands or specific management regimes applied. Clearcutting, however, is the standard practice but subsequent silvicultural practices vary (Nova Scotia DNR 2000). The concentration of wind disturbed forests in the older age range reflects the fact that wind disturbance in Nova Scotia is generally of low intensity, allowing trees to be- come much older than does short-rotation forestry disturbance. Managed forests were common and thus chosen randomly from a larger subset of forests; the rarity of naturally disturbed spruce forests, on the other hand, allowed no room for random selection. Table 1 shows the distribution of the study forests among disturbance history classes and age classes. It is evident that a rigorous sampling design with regard to stand age and distur- bance was not possible to achieve and has been addressed in the analysis (see Analysis). The remaining three forests were fire origin and dominated by black spruce (Picea mariana), a species dependent on fire for regeneration when not in lowland or bog soils (Fowells 1965). Black spruce and red spruce are known to hybridize and they are often difficult or impossible to distinguish (Roland and Smith 1969). Forest habitat structures.In each study forest, deadwood volume and crown clo- sure were sub-sampled in six 15 x 15 m plots located at 40 m intervals along two parallel transects spaced 40 m apart. For each piece of deadwood, the length and diameter at each end were measured (or estimated) to calculate volume based on the frustum of a cone. Tree species and decay states (see Table 2) were recorded for each piece of deadwood to allow independent tallying of deadwood in different classes. To Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 313 Table |. Distribution of forests delineated by age and disturbance history classes. Disturbance History Class Codes for Mean Canopy Tree Age Classes Reference 30-49 years | 50-69 years | 70 years + clearcut origin CLEARCUT 1 6 4 clearcut and subsequently |CC&THINNED 6 2 0 thinned naturally disturbed FIRE I vr 1 (fire origin) naturally disturbed WIND 1 2 5 (wind only) total 9 11 10 estimate percent crown closure, two methods were used. In most forests, a flat mir- ror with 20 equally spaced dots was held horizontal at chest level to determine the proportion of dots occurring in crown-covered space. In remaining forests, a short cardboard tube was used as a scope to visually estimate proportions of crown closure. Applying both methods in one forest to test comparability returned similar results. Three crown closure readings were taken at random distances (10-20 m) from each of the six plot centers through late July to early August (during peak deciduous foliage). Within each plot, a healthy dominant canopy tree was cored to estimate tree age for a total of six trees per forest. Descriptions of all variables measured are given in Table 2. Beetle sampling and identification.1o sample saproxylic beetles systematically at each forest, a window flight intercept trap (FIT) design was employed. The FIT is passive in that it does not attract beetles, thus providing random samples of the fauna that actively fly in the local environment. @kland (1996) showed that window FITs were effective for providing large samples of saproxylic beetles and Wikars et al. (2005) showed that window FITs compared with other methods caught the greatest number of species. In a single season Muona (1999) found that FITs collected 50.6% of the species of beetles (and 48.3% of forest species) in a study in the Oulanka region of Finland. Emergence traps where beetles are sampled as they emerge from experimental logs (Gibb et al. 2006) are logistically difficult. This was not a popular method for sam- pling saproxylic beetles in the field when this study was conducted in 1997 and thus, was not considered an option for this study. We employed a design consisting of two bisecting 30 x 30 cm transparent plastic (Lexan 0.030) panes, covered with a white plastic (Styrene 0.040) conical roof for rain shelter, and attached to a Styrene collecting funnel below. Removable plastic sample jars attached at the base of the collecting funnel held a 50 % diluted ethylene glycol killing/preserving solution. In each forest site, six FITs were hung from existing tree branches, with the bottom of the trapping surface approximately 1 m above the forest floor. FITs were located 40 m apart along two parallel transects separated by 40 m. All FITs were set up during 13-19 May 1997 and run until 13-15 August 1997. Catches were collected every two weeks within 3 days of each other, and the ethylene 314 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Table 2. Description (A) forest structure variables measured in each forest, (B) deadwood diversity and (C) beetle diversity attributes. Variable Name Description (A) CROWN crown closure (%) SIZEL volume of deadwood with diameter 0-15 cm (m? /100m7) SIZE2 volume of deadwood with diameter 16—30 cm (m? /100m/7) SIZE3 volume of deadwood with diameter >30 cm (m? /100m7) DECAY1 volume of deadwood with little or no decay (m* /100m/’) DECAY2 volume of deadwood with 1—5 cm decay depth (m* /100m‘) DECAY3 volume of deadwood with >5 cm decay depth (m° /100m7) STANDING volume of deadwood standing, > 45° from floor (m? /100m’) FALLEN volume of deadwood fallen, < 45° from floor (m?* /100m7) DECIDUOUS volume of deadwood in all deciduous species (m° /100m”) CONIFER volume of deadwood in all coniferous species (m? /100m”) ALLDWOOD volume of all deadwood measured (m? /100m/7) (the summation of deciduous and coniferous deadwood) (B) SHAN-SIZE Shannon diversity index of deadwood in 3 size classes SHAN-DECAY Shannon diversity index of deadwood in 3 decay classes SHAN-POS Shannon diversity index of deadwood in 2 position classes (standing and fallen) SHAN-D/C Shannon diversity index of deadwood in 2 tree type classes (deciduous or conifer) SHAN-ALL Shannon diversity index of all 10 deadwood classes (above) (C) ABUNDANCE total number of beetles from all species sampled RICHNESS total number of beetle species sampled RICH/ABUN richness/abundance of all beetles in each site SHANNON Shannon diversity index of all beetles in each site RARE-1 number of species sampled at site for which the total catch was | individual RARE-5 number of species sampled at site for which the total catch was 5 or fewer RARE-10 number of species sampled at site for which the total catch was 10 or fewer RARE-15 number of species sampled at site for which the total catch was 15 or fewer RARE-30 number of species sampled at site for which the total catch was 30 or fewer RARE-60 number of species sampled at site for which the total catch was 60 or fewer glycol solution was changed at this time. This gave six two-week samples for each trap, with the collection dates on or adjacent to 3 June, 16 June, 1 July, 13 July, 30 July, and 14 August. Beetles were separated from each trap sample within one week and preserved in 70% acetic ethyl alcohol. Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 315 All beetles were identified to species although only species considered to be saproxy- lic were employed in subsequent analyses. Saproxylic (sensu lato) is defined here in ac- cordance with Speight (1989). The inclusion of species in this category was made on a specific or, more commonly, generic basis, consulting a wide variety of published sources (commencing with Arnett and Thomas (2000) and Arnett et al. (2002), fol- lowed by family-specific treatments), or if such information was not readily available in the literature, consulting with specialists of different Coleoptera families listed in the acknowledgments. Appendix 1 gives the full list of saproxylic species caught. Species identifications were initially made by D.J. Bishop employing the reference collection of the Canadian National Collection of Insects, Arachnids, and Nematodes. Subsequently the process of identification was continued by C.G. Majka and employing the reference collection of the Nova Scotia Museum. At both stages the process was great- ly assisted by Coleoptera specialists (listed in the acknowledgments) who were able to identify difficult species or confirm determinations done by Bishop or Majka. The general systematics and taxonomy follow Arnett and Thomas (2000) and Arnett et al. (2002). Statistical analyses. The aim of the initial analysis was to test whether habitat structures differed according to the four disturbance history classes: (1) clearcut, (2) thinned, (3) fire, and (4) wind. For this, distribution-free Kruskal-Wallis tests (non- parametric analogs to one way analysis of variance) were used, followed by the Nu- menyi a posteriori multiple comparison tests in cases where the initial tests suggested a significant (p<0.05) difference (described in Zar 1999). To test predictions at the alpha scale, the initial procedure was to reduce the structural information and the beetle assemblage from each forest to meaningful diversity attributes. The Shannon diversity index (Brower et al. 1990) was selected to index both entities. The Shannon diversity value was calculated independently for four aspects of dead- wood habitat diversity for each forest: (1) decay classes, (2) size classes, (3) standing/ fallen classes, and (4) conifer/deciduous classes. The Shannon diversity for measures of beetle assemblage was calculated without refinement (Table 2). The habitat diversity and beetle assemblage attributes were independently tested for any differences with respect to disturbance history using parametric, one-way ANOVA tests, with Tukey HSD multiple comparison post hoc tests to detect the location of any significant (p<0.05) differences between any two disturbance history classes. Taking the measures of the beetle assemblage to be the dependent variables and the forest habi- tat diversity variables as the independent variables, stepwise multiple linear regression analysis (Weisberg 2005) was used to test if and how aspects of the beetle assemblage were related to deadwood habitat heterogeneity. Assumptions of independence and homogeneity of variance were checked with plots of residuals. The assumption of nor- mality of residuals was checked with histograms of residuals for each test. Analysis at the beta scale comparing beetle community turnover and deadwood structure turnover among forest sites in relation to disturbance history class was carried out by non-metric multidimensional scaling (NMS) (see Clark 1993) and SIMPER and ANOSIM analyses in Primer 5 2002 (PRIMER-E Ltd. 6 Hedingham Gardens, Roborough, Plymouth PL6 7DX, United Kingdom). 316 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Secondly, multi-response permutation procedures (MRPP) were used to test whether beetle- and habitat-defined forests differed across disturbance history class- es. This non-parametric analog to discriminate analysis circumvents assumptions of normality and homogeneity of variance that could not be met with the current data set. Ihe MRPP method also returns the mean distance among sample points within each disturbance history class, which describes the relative heterogeneity among for- ests in each class. For the distance measure in MRPP, we used the Sgrenson coeflicient (also known as the Czekanowski or Bray-Curtis coefficient). It measures percent dissimilarity (PD) between two samples, calculated as PD = 1—2W/A+B, where W is the sum of shared abundances and A and B are the sums of abundances in individual sample units. For analysis at the gamma scale, species-sample curves were constructed to test the prediction that cumulative species richness for all naturally disturbed forests (fire and wind disturbed forests inclusive) was higher than that for all forests under even-age management (clearcut and thinned forests inclusive). Species-sample curves were also constructed individually for each disturbance history class to allow a visual evaluation of the gamma richness trends for each. Statistical analyses were carried out with SYSTAT Version 5.02 (Systat Inc. 1993), S-PLUS (MathSoft 1999)and PC-ORD Version 3.09 (McCune and Mefford 1997). Results A total of 12,151 beetles of 389 species were collected. Of these, 296 species compris- ing 10,488 specimens in 45 families were determined to be saproxylic species (Appen- dix 1). Two species, Anapsis rufa Say and lsomira quadristriata (Cooper) were caught at all 30 sites and were the most abundant with 3,880 and 1,129 specimens respectively. Eighty-five species were represented by single captures and 216 species were repre- sented by less than 10 specimens. Forest structure generally showed high variation within disturbance history classes. Significant disturbance-class differences tested by a Kruskal-Wallis analysis were identi- fied for 2 (CROWN 247=10.54, p= 0.015, SIZE1 X7=8.36, p=0.039, and marginally DECAY? X’=6.34, p=0.07) of the 12 variables tested. Crown closure (CROWN) was highest in wind disturbed and clearcut forests and lowest in fire disturbed forests. Small-diameter deadwood (SIZE1) showed significantly higher volumes in thinned forests than in either clearcut or wind disturbed forests. DECAY2 was greatest in WIND disturbed sites. Stand age was not a function of any of the deadwood measures among stands. Alpha diversity. The ANOVA analysis to test whether naturally disturbed forests exhibited higher diversity of deadwood structures offered evidence for one of the five attributes tested. SHAN-DECAY (diversity of deadwood decay classes in each forest) was significantly higher in both fire (p=0.040) and wind (p=0.004) disturbed forests than in thinned forests (F=4.75, p=0.01). Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 317 The general linear regression models constructed to determine whether the beetle diversity could be explained by habitat diversity attributes with disturbance class and age as factors resulted in four models (Table 3). In interpreting these results, it should be borne in mind that the dependant variable is a function of the explanatory variables. If the coefficients are negative, the depend- ant variable is a negative function of the explanatory variable, and if positive a positive function. The F statistic represents the full model and the T/P are the univariate results. Beta diversity. [he forest sites plotted in the space of the beetle assemblage show two important patterns: differences in the inter-site distances within disturbance classes and actual segregation of forest disturbance classes (Figure 1, Table 4). Clearcut forests show the tight- est aggregation (mean distance = 0.421), followed by fire disturbed forests (mean distance = 0.571), then thinned forests (mean distance = 0.581), with wind disturbed forests showing the greatest heterogeneity (mean distance = 0.603) which was statistically significant (Table 4). Different beta patterns were observed in forests with regard to habitat structures. Figure 1 shows that, unlike the species-defined plot, fire disturbed forests show the Table 3. Significant multiple linear regression models of beetle assemblage attributes (dependent vari- ables) regressed against habitat structure diversity attributes (explanatory variables). Construction by for- ward and backward stepwise inclusion and exclusion of variables, based on two-tailed p-value of 0.1. T is the test statistic; P is the probability value. Dependent | Explanatory | Coefficient T/P F- | Model | Multiple Variable Variable(s) +SE (2-tailed) |statistic| P-Value R? ABUNDANCE Intercept -216.744+163.13 | -1.33 / 0.20 0.01 0.29 AGE 1.28+0.97 1324-020 DISTAGEASS |" 1-0.72236.692) |"O2951 0.47 SIZE! 1778.44+574.00 | 3.01 / 0.005 SHAN-DIC_ | 584.43+186.75 RICH/ABUN Intercept 0.28+0.04 6.87 / 0.00 ; 0.005 0.37 AGE 0.00+0.00 -1.85 / 0.08 DIST. CLASS 0.01+0.01 O20 32 SIZE1 -0.61+0.17 | -3.48 / 0.00 SIZE2 0.33+0.14 22 F-10039 DECIDUOUS SHANNON Intercept 3.82+0.40 9.65 / 0.00 0.03 0.24 AGE -0.0040.00 | -1.94/ 0.06 DISHCTASS 0.07+0.09 0.81 / 0.42 SIZE1 -4.3641.39 | -3.13 / 0.00 SHAN-D/C -1.00+0.45 | -2.21 / 0.04 RARE-10 Intercept 68.96+10.89 | 6.35 / 0.00 : 0.10 0.10 AGE 0.05+0.05 1.00 / 0.33 DIST. GLASS -0.03+1.81 | -0.02 / 0.99 CROWN -27.91+13.44 |-2.08 / 0.05 DECIDUOUS] 47.62+22.90 | 2.08 / 0.05 318 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Table 4. Results of multi-response permutation procedures (MRPP) analysis, testing for separa- tion of disturbance history classes based on (A) beetle assemblage and (B) forest habitat struc- tures. Mean distance within class can be read as a relative measure of beta heterogeneity within disturbance history classes. P-values associated with multiple comparisons denote significance of group separation. Adjusted multiple comparison a level for p-value significance at 0.05 = 0.009, and for significance at 0.10 = 0.017, based on adjusted a = 1-(1-c)!/ umber oF comparisons CLEARCUT | CC-THINNED | FIRE | WIND | All-groups p-value (A) mean distance within class 0.421 0.581 0.571} 0.603 CC&THINNED 0.027 FIRE 0.001 0.075 0.0006 WIND 0.005 0.062 0.025 (B) mean distance within class 0.217 0.296 05353-| 30.175 CC&THINNED 0.219 FIRE 0.246 0.754 0.097 WIND 0.147 0.060 0.078 Forests in space of 294 species t20 fle [] ig c6 Disturbance History Classes * a Oo fire origin predominantly wind clearcut clearcut and thinned Figure |. Non-metric multidimensional scaling ordination diagrams of forest sites in two-dimensional space defined by (A) beetle assemblage and (B) habitat structures. Symbols signify disturbance history class of forests: A= CLEARCUT, O= THINNED, M= FIRE, and @= WIND. Numbers identify the specific forest. Forest number 25 was excluded as an outlier in (B) due to much higher deadwood volumes than all other sites. Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 319 greatest heterogeneity (mean distance = 0.353), followed by thinned forests (mean dis- tance = 0.296) and then clearcut forests (mean distance = 0.217), and wind disturbed forests with the lowest heterogeneity (mean distance = 0.175) but the statistical differ- ence was only significant at the 0.1 probability level (Table 4B). However, one must note that an outlying wind disturbed site (number 25) had an extremely high volume of deadwood and was removed due to difficulty in interpreting the NMS diagram oth- erwise. Ihe MRPP for wind-disturbed forests, when the outlier was included, ranked the wind-disturbed class as second most heterogeneous, behind fire-disturbed forests. The ANOSIM analysis testing for pair-wise similarities among disturbance classes gave a global R statistic of 0.174 with p=0.09 but none of the pair-wise R values were significant even at p=0.10. Results of the SIMPER analysis on average dissimilarity among pairs of sites by disturbance are presented in Appendix 2. Gamma diversity.Of the 296 native saproxylic species sampled, 227 (77.2%) were sampled in the naturally disturbed forests and 228 (77.6%) were sampled in the man- aged forests. The first and second order Jackknife estimates of total species richness were 301.7 and 337.2 for naturally disturbed forests and 308.3 and 351.4 for man- aged forests. This does not support the prediction that the total fauna richness would be higher in naturally disturbed forests; in fact, the opposite appears more likely to be true. The species-sample curves constructed for the four disturbance classes (Figure 2) SPECIES-SAMPLE CURVES 220 + species I 2 3 4 5 6 7 8 9 10 i samples Figure 2. Species-sample curves for each forest disturbance class. 320 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) suggest the following relationship of species richness across disturbance history classes: thinned>wind>fire>clearcut. Discussion 4.1 Alpha diversity. ‘The analysis of deadwood structure and diversity at the alpha level revealed two insights about thinned forests: there is both a higher volume of small-di- ameter deadwood and a lower decay-class diversity in thinned forests compared to wind and fire disturbed forests. Both pre-commercial and commercial thinning treatments were represented in the thinned forest class (five and three forests, respectively). While thinning treatments are applied to stands of varying ages, the results are typically an im- mediate increase in the volume of similar-diameter downed deadwood. One would pre- dict that this would lead to local homogeneity of deadwood size-structure, though this finding was not supported in our results. The senescence of the downed deadwood in a single cohort predictably results in decreased decay variation in space, as supported by the lower decay-class diversity in thinned forests. The higher volume of small-diameter deadwood in thinned forests probably reflects the dominance of pre-commercial thin- ning in the thinned class, because pre-commercial treatments are applied in forests with young, small trees. Thus, our results suggest that lower diversity and increased volumes of deadwood may be generally associated with the thinning practice. Saproxylic beetle abundance was a positive function of small diameter deadwood volume and the diversity of total deadwood volume. Diversity of saproxylic beetles measured as richness divided by abundance, however, was a negative function of small diameter deadwood and deciduous deadwood but a positive function of intermediate size deadwood. Shannon diversity of saproxylic beetles was a negative function of the diversity deadwood (deciduous or coniferous) volume. Thus the relationships are not clear. Rare beetles were a negative function of crown closure and a positive function of deciduous deadwood volume. One post hoc observation is worthy of mention: deadwood diversity levels in the managed forests might largely be an artifact of pre-harvest forest environments. Nearly all managed forests in this study were in their first or second even-age rotations. Hence, most of these forests likely contained large amounts of residual deadwood from decid- uous species and large diameter trees, patterns one would not expect to endure through continued successive short-term rotations. Since second growth forests will usually be harvested before they can reproduce their own old-forest characteristic structures, we can only anticipate their eventual disappearance. In so much as these decomposing artifacts are microhabitats and resources, we can of course also anticipate the disap- pearance of the species that depend on them. One explanation for the lack of clear results may be that deadwood may not have been measured over an area that was adequate to reflect any deterministic influences on local beetle assemblages. One cannot necessarily assume that a sampled array of fly- ing beetles will reflect only the microhabitats available within a few hundred meters of Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 321 the sample points. In theory, the deadwood sampling area that is ideal for discerning relationships emerges from a combination of ‘best’ areas for individual species and in- dividual forests. In recognizing this quandary of FIT sampling, Okland (1996) tested the habitat relationships for individual beetle species in Norway and found that habitat measurements collected over an area of 32 hectares produced the highest number of re- lationships. To assume that the measurements in this study represented the surround- ing forest environment over this large an area may be unreasonable. In a study comparing FIT sampling to bark-peeling as methods of determining saproxylic faunas, Siitonen (1994) found that the numbers of species and individuals of aerially-dispersing beetles captured by FITs were unaffected by the quantities of deadwood in the local area of the trap. He interpreted this to mean that the distribu- tion of such aerially dispersing species was related to larger forest scales and not the immediate local conditions. Since the present study employed FITs, the beetle assem- blages sampled may have been reflective of such larger forest scales. Gibb et al. (2006) found that saproxylic insects trapped using emergence traps showed a poor relation- ship with deadwood at local scales. Thus, in the future a combination of FIT and emer- gence trapping should probably be employed and larger stands, if available, should be sampled. Other techniques such as baited Lindgren funnel traps, bark peeling, car nets, hand collecting, and light-trapping may also yield beetles, particularly rare species, not sampled by conventional techniques (Muona 1999). Beta diversity. While the alpha and gamma analyses focus on the unqualified di- versities of habitats and beetle assemblages, within forests and within disturbance class- es, the beta analyses examine the relative composition of the beetle assemblages and deadwood structures. The beta analyses revealed that actual species compositions of the assemblages were not clearly different among classes, and that higher turnover among naturally disturbed forests was evident. Observations of high turnover of saproxylic beetle species without changes in species richness were made in central Finland by Kaila et al. (1997) when comparing recent clearcuts with mature forests; by Vaisanen et al. (1993) when comparing the fauna of birch trunks in old-growth forests with those in managed forests; and by Sippola et al. (2002) in comparing saproxylic beetle diversity in old growth and clearcut regeneration stands in Finnish Lapland. While the current study does not support the prediction of reduced species diversity related to reduced deadwood habitat diversity in managed forests, it does support the claim that managed forests support different faunal assemblages (Niemela 1997). The larger heterogeneity of fire disturbed forests is in keeping with studies that indicate that many saproxylic species are adapted to fire disturbances and subsequent forest succes- sion (Granstr6m 2001; Buddle et al. 2006). Siitonen (2003) identified the virtual elimina- tion of fire disturbances in Fennoscandian boreal forests as one of two principal conservation concerns for saproxylic species (the other being the low proportion of old growth forests). Similarly, in relation to wind-disturbed forests, Duelli et al. (2002) documented dramatic short-term (over the span of a decade) increases in the species richness and biodiversity of invertebrates, reptiles, and small mammals in severe windthrow sites in Switzerland follow- ing a 1990 storm. At these same sites Wermelinger et al. (2002) found profound changes 322 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) in saproxylic beetles with certain groups (Cerambycidae and Buprestidae) becoming 30 to 500 times more abundant than in adjacent intact forests. Although the dramatic increases in numbers of individuals were relatively short-lived, the composition of the saproxylic beetle fauna became progressively more dissimilar from the control plots. Gamma diversity. The final prediction of lower gamma diversity of saproxylic bee- tle species in managed forests compared to naturally disturbed forests was not sup- ported. The prediction was partly underpinned by an assumption that forest habi- tat structures differed between managed and naturally disturbed forests at the alpha scale, which was not highly supported. If anything, the higher species-sample curve in thinned forests seems to suggest that thinning disturbance generates more species than either wind of fire disturbance. Otherwise, the non-thinned clearcut forests appeared to have a lower diversity of species than both wind and fire origin forests. In Swedish forests Larsson et al. (2006) found that ‘new forestry’ management prac- tices that left large volumes of deadwood in stands improve conditions for some species of saproxylic beetles. Furthermore saproxylic beetles as a group are a composite, reflective of different stages of wood decay. Sippola et al. (2002) examined saproxylic and non-saprox- ylic beetle diversity in old-growth and regeneration forests in Finland. Although no sig- nificant differences were detected in the rarefied number of species between old-growth and regeneration stands, the species composition of both saproxylic and non-saproxylic beetles differed. Species colonizing recently dead trees, soil-dwelling open-habitat species, and some polypore-dwelling ciids were more abundant in recently cut and regenerat- ing sites, whereas many mycetophagous beetle families were almost completely absent. Overall saproxylic species richness or even species diversity does not discriminate between these different saproxylic components and may mask important differences between for- est beetle communities in forest stands of different age, composition, or stand history. Conservation requirements. [he general claim that modern forestry in northern forests has altered invertebrates by altering forest habitat structure is well supported in the literature (Niemela, 1997; Martikainen et al. 1999, 2000; Siitonen 2001; Grove 2002; Jonsson et al. 2005). In Nova Scotia, Dollin et al. (2008) found that harvested forest stands had lower Coleoptera species richness and were host to a significantly different suite of species than unharvested stands. Two tentative conclusions from the current study help extend this claim in Nova Scotia forests: 1) the species assemblage of saproxylic beetle species differed between managed and naturally disturbed forests, and 2) these beetle assemblages are more variable among naturally disturbed forests than among managed forests. If management is to decrease the altering effect of forestry prac- tices, the next requirement is to identify or confirm the particular aspects of forest struc- ture that are being altered and that are in turn altering biodiversity (Essen et al. 1992; Swanson and Franklin 1992; Noss and Cooperrider 1994; Franklin 1995; Lubchenco 1995; Niemela 1997; Siitonen 2001, 2003; Grove 2002; Langor et al. 2006). Research from countries with longer histories of intensive forestry has revealed sev- eral cases of invertebrate endangerment due to loss of microhabitats found in deadwood or moribund trees. Some of these microhabitats are lost because they are found in tree species that are not economically desirable, like aspen (Siitonen and Martikainen 1994) Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 323 and beech (Nilsson and Baranowski 1997). Others are threatened because they are part of trees with large diameters, which are not maintained amid short-lasting even-age stand rotations (Okland et al. 1995; Nilsson et al. 1995; Siitonen and Saaristo 2000). These and other factors have lead to concern about the diversity of deadwood microhab- itats in general, as defined by decay states, sizes, and tree species (Okland et al. 1995). If the goal in managing forests in Nova Scotia is to maintain patterns of heteroge- neity similar to those that arise after natural disturbances, the indication offered here by saproxylic beetles is that this goal is not being met. Majka (2007) examined 14 families, subfamilies, and tribes of saproxylic beetles in the Maritime Provinces of Can- ada and found 59 apparently rare species (representing < 0.005% of specimens from the region) that comprise 33% of the 178 species within these groups. Majka (2007) proposed that this apparent scarcity of a large proportion of the saproxylic fauna might be due to the history of forest management practices in the region. If Nova Scotia has not yet suffered biodepletion to the extent of European forests, this may only reflect the fact that not enough time has passed for the deadwood lingering from old-forests in second-growth forests to fully return to soil. Acknowledgments R. Cameron assisted the forest selection process. L. Parriag assisted field sampling and laboratory work. The following coleopterists kindly assisted with identifications: J. Cook (Carleton University, Ottawa), E Andrews and A. Cline (California Department of Food and Agriculture), D. Chandler (University of New Hampshire), J. Klimaszewski (Cana- dian Forest Service), D.B. McCorquodale (Cape Breton University), M. Sérensson (Uni- versity of Lund, Sweden), M. Thayer (Field Museum of Natural History, Chicago), W. Ricker (Latridiidae.com), Y. Bousquet, D. Bright, A. 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Disturbance Classes Wind disturbance 8 stands Clearcut | Clearcut + | Fire origin 11 stands thin 4 stands 8 stands Trophic Category Aederidae Aderinae Vanonus wickhami Casey | Xylo | 0 | Fl! -O50) iP a | 0.5 | 1 | 0.1 | 1 Anobiidae Anobiinae Hadrobergmus notatus (Say) Hemicoelus carinatus (Say) Microbregma e. emarginatum (Duftschmid) + Platybregmus canadensis Fisher Dorcatominae Caenocara oculata (Say) Dorcatoma falli White Dorcatoma pallicornis LeConte Sculptotheca puberula (LeConte) Xylo 0 | 05) 0 0 0 | 0 | 0.3 Ptiliinae Ptilinus ruficornis Say Xylo 0 | 0) 0.4 | 2 0 0 0 0 Buprestidae Buprestidae Melanophila d. drummundi (Kirby) Xylo 0g) 6) 20z)) Dsl EL 0 0 0 0 Carabidae Harpaliinae Bradycellus nigriceps LeConte Bradycellus nigrinus (Dejean) Bradycellus semipubescens (Dejean) Cymindis limbatus Dejean Dromius piceus Lindroth Syntomus americanus (Dejean) Trechinae Bembidion mimus Hayward Pred CBRN Res CB | es a 0 0 0 0 Trechus rubens (Fabricius) + Pred O21e5) Pl 0 0 0 0 0 0 Cerambycidae Aseminae Asemum striatum (Linnaeus) Xylo 0 Crash 0 0 0 0 Tetropium cinnamopterum (Kirby) Xylo OFS el 0 0 0 0 0 0 Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 329 Disturbance Classes Trophic Clearcut Clearcut + | Fire origin . Wind Cateeocy 11 stands thin 4 stands | disturbance 8 stands 8 stands Cerambycinae Molorchus b. bimaculatus Say Sees | || 0? 80" 0.08 | 40 |) oor oe | 0 Phymatodes testaceus (Linnaeus) t Xylo O21 | ad 0 0 0 07 || <0: 1 Laminae Hyperplatys maculata Haldeman Xylo Oe | 20> |.470 0 OF IFO Oatles |e Monochamus s. scutellatus (Say) fol o | 0 Pogonocherus penicillatus LeConte 0 Lepturinae Acmaeops p. proteus (Kirby) Xylo 0 | 0 0 Oe FEOES. || Pel 0 0 Acmaeopsoides rufulus (Haldeman) Xylo 0 |0]}] 0 OF | SOF Ml 0 0 Anthophylax attenuatus (Haldeman) Xylo 0> [| 05) 0 0 e6nml Eos] oe24| 2 Anthophylax cyaneus (Haldeman) Xylo 0° |20"| 0 0 Ow nO” || 0725 |e Anthophylax viridis LeConte Xylo Qaln| Pl? |) 0568 | 3/805 || ele O39 | Be Evodinus m. monticola (Randall) Xylo 34 | | MeleOe PO Z a be 9 Idiopidonia pedalis (LeConte) An 2d ® || 25, Pores O45 4 Judolia m. montivagens (Couper) Selo | 0: MeO Oa) al | toe [Ex a 3 Leptura subhamata Randall 1 Os | 2D |eo. | Ore: |b Lepturobosca chrysocoma (Kirby) 0 1) WP s03301) 20 0 0 Pidonia rufficollis (Say) i 4 S| P28 || ae Le a 5 Stictoleptura c. canadensis (Olivier) 1 Strangalepta abbreviata (Germar) 8 Trachysida aspera brevifrons (Howden) | _ Xylo Oise | Sl 0 Os 820.3% |-Al 0 0 Trachysida mutabilis (Newman) Xylo 0 ad Poa Os i Pd 0 0 0 0 Xestoleptura tibialis (LeConte) Xylo 0 0 0 0 0 0} 0.1 1 Cerylonidae Cerylonidae Cerylon castaneum Say Ciidae Ciidae Ceracis sallei (Mellié) Ceracis thoracicornis (Ziegler) Bolito Cis fuscipes Mellié Bolito Cis horridulus Casey Orthocis punctatus (Mellié) Clambidae Clambus pubescens Redtenbacher * Rea 36 2 P Myceto Cleridae Clerinae Epiphloeinae Madoniella dislocata (Say) 1 330 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Disturbance Classes Trophic Clearcut | Clearcut + | Fire origin Wind ee P 11 stands thin 4 stands | disturbance oe 8 stands 8 stands Av.S|_S Corylophidae Orthoperinae Orthoperus suturalis LeConte Wiyceto- | -Oels| als) S051 |. La) e0S-is) fel 0 0 Pityophthorus opaculus LeConte Xylo 0; O 0 1 2s | , ih if Ampedus luctuosus (LeConte) Xylo 0 0 | «OS | 3 Ampedus mixtus (Herbst) Xylo Ori Ply || es. | 2 Ampedus protervus (LeConte) Xylo Oriee| ee Oaks |L.t Ampedus pullus Germat | Xylo | 0 |o} o [0 | Ampedus semicinctus (Randall) SSeloe | G0: seo OF |e" Ampedus sp. undescribed Dalopius cognatus Brown Dalopius fuscipes Brown Dalopius gentilis Brown 0.1 0.1 WN} | Oo -) KSBLOPAIN IAI Oslo; Re ;os;o]& 0 2 1 1 1 4 i=) Oo o};|o =) \9 o};|o i) Oo Datlopius pennsylvanicus Brown | Myceto | 0 | 0] 01 | 1 | Datlopius vagus Brown Myceto | 4.6 | 2 | 4.3 0.1 Prosterninae Athous brightwelli (Kirby) Pred Ale) | ol 0 0 0 2 Athous orvus Becker | Pred | 0 | o|] o [0] 2 Athous rufifrons (Randall) 6 rh 2 8 Athous scapularis (Say) 0 0 0 1 Beckerus appressus (Randall) 1 0 0 0 Denticollis denticornis (Kirby) , 6 : 8 3 9 Eanus maculipennis (LeConte) | Pred | 0 | 0] 08 | 2 | For! 0 Hypoganus sulcicollis (Say) Pred 0/0]; O 0 Liotrichus falsificus (LeConte) Pred 293) 72 |e Peer | Ke OB gear | 1.8 Liotrichus spinosus (LeConte) Pred oS ee ane: Am 25 8 Liotrichus vulneratus (LeConte) | Pred | 0 | Or OSs eZ $i Nitidolimonius resplendens (Randall) 23 Ose, Eat a Metanomus insidiosus (LeConte) 5 6 | We3? || 28| Uhl 5 Oxygonus montanus Schaeffer 3 3 zie! Oa), telnm | aa) Pseudanostirus hamatus (Say) 0 1 0 Pseudanostirus p. propolus (LeConte) 0 0-8-0. 3ail| le O55 15 Pseudanostirus triundulatus (Randall) Se a Raed bear co PD 332 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Disturbance Classes Trophic Clearcut | Clearcut + | Fire origin Wind Cae P 11 stands thin 4 stands | disturbance ahora 8 stands 8 stands Endomychidae Epipocinae Hadromychus chandleri B & L Lycoperdininae Mycetina perpulchra (Newman) 03/1] o Jo} o Jo] o [o Myceto™ | 0:4 |S |G:0s60%) “4% |> O85. (ll | 20.2. |e Merophysinae Phymaphora pulchella Newman Erotylidae Tritominae Myceto | 0.3 | 2 | 0.5 | 3 0 0 | 0.1 1 Triplax dissimulator (Crotch) Bolito 0 0 0 Gy | 0; = 1 0 0 Triplax flavicollis Lacordaire Tritoma pulchra Say Eucinetidae Eucinetus morio LeConte Bolito 0 0 0 0.1 1 Nycteus punctulatus (LeConte) Eicnemidae Melasinae Epiphanis cornutus Eschscholtz Hylis terminalis (LeConte) lsorhipis obliqua (Say) Microrhagus pectinatus (LeConte) Histeridae Dendrophilinae Paromatus teres LeConte | Pred | 0.1 | 1 | 03 | 2 | Lo | o1 | 1 Laemophloeidae Laemophloeinae Laemophloeus fasciatus Melsheimer__| Myceto | 0 |0| 0 |0]} 0 [0] 01 Lampyridae Lampyrinae eye corrusca e{hintiaeus) Pred 3 10 Lace ana Olives) [Ped 0 [oor [1 | 0 [olor Pyractomena angulata (Say) |_Pred_| 0.1 | 1] 05 | 1] 05 | 1] 01 | 1 Latridiidae Cortcariinae Corticaria speciés Myceto | 0. 0:3. | 2 Grn goa HetE | wesw [oa Tos Ts 1 fT Melanophthalma americana Myceto | 0. 6 0.6 10 QF | > Mannerheim Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 333 Melanophthalma inermis Motshulsky Melanophthalma pumila (LeConte) Latridiinae Disturbance Classes Trophic Clearcut | Clearcut + | Fire origin Wind P 11 stands thin 4 stands | disturbance 8 stands 8 stands Av. S |} o |o} o | Cartodere constricta (Gyllenhal) + Enicmus tenuicornis LeConte Latridius minutus (Linnaeus) t Stephostethus liratus (LeConte) Leiodidae Leiodinae Agathidium difformis (LeConte) Agathidium fawcettae Miller & Wheeler Bolito 0 0] 0.4 | 2 0 Anisotoma basalis (LeConte) Anisotoma blanchardi (Horn) Anisotoma geminata (Horn) Bolito 73 Bolito 0 : | Bolito | 0 | 0] 0 | 0| | Bolito | 0 |o| 0 | 0| Anisotoma horni Wheeler Bolito Anisotoma inops Brown Bolito Leiodes assimilis (LeConte) Bolito Lucanidae Syndescinae Ceruchus piceus (Weber) | Xylo Gel et MPS 2 ae | 0.3 | 1 | 0.2 | 2 Lycidae Erotinae Dictyopterus aurora (Herbst) Eros humeralis (Fabricius) Myceto | 0.4 | 2] 0.4 | 2] 0.5 | 2 | 03 | 3 Lycinae Leptoceletes basalis LeConte Platoderinae Plateros lictor (Newman) Plateros subfurcatus Green Myceto | 0.1 | 1 | 0.1 | 1 | 0.3 | 1 | 0.4 | 4 0.4 0 0 0 0 0 Myceto | 0.1 | 1 0 Melandryidae Melandryinae Dircaea liturata (LeConte) Emmesa connectans Newman Hypulus simulator (Newman) Orchesia castanea (Melsheimer) Prothalpia undata LeConte Scotochroa atra LeConte Scotochroa buprestoides (Kirby) Scotochroides antennatus Mank Serropalpus substriatus Haldeman N PW} OP RN RFR LO [Nn 334 Spilotus quadripustulatus (Melsheimer) Symphora flavicollis (Haldman) Xylita laevigata (Hellenius) DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Disturbance Classes Trophic Clearcut | Clearcut + | Fire origin Wind ae P 11 stands thin 4 stands | disturbance ahora 8 stands 8 stands | Xylo | o fo] o jo} o |o] o1 | o1}1} o [o] o |o| 02 | Xylita livida (Sahlberg) * Monotomidae Rhizophaginae Mycetophagidae Mycetophaginae Mycetophagus pluripunctatus LeConte Nitidulidae Carpophilinae Myceto 0 Carpophilus brachypterus (Say) | sap | cor [se"] or eos] ws) [ae ho "eo Cillaeinae Colopterus truncatus (Randall) Cryptarchinae Glischrochilus fasciatus (Olivier) | Sap | o folor}il} o jojo) i Glischrochilus sanguinolentus (Olivier) Glischrochilus siepmanni Brown Epuraeinae Epuraea aestiva (Linnaeus) Epuraea erichsonii Reitter Epuraea labilis Erichson Epuraea rufida (Melsheimer) Epuraea terminalis Mannerheim Epuraea truncatella Mannerheim Oedemeridae Calopodinae Calopus angustus LeConte Pyrochroidae Pyrochroinae 0 Dendroides canadensis Latreille 0.3 Dendroides concolor (Newman) Schizotus cervicalis Newman Pythidae Priognathus monilicornis (Randall) 0.6 Salpingidae Salpinginae Scarabaeidae Salpingus viridiaeneusRandall___—_|_ Xylo_ | 0.1] 1] 01 |1] 0 Jo} o | 0 Cetoniinae Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia Trichiotinus assimilis (Kirby) Scirtidae Cyphon collaris (Guérin-Meéneville) Disturbance Classes aie Trophic Clearcut | Clearcut + | Fire origin Wind ae P 11 stands thin 4 stands | disturbance arena 8 stands 8 stands Av. S | Xylo | o Lo] o Jol o Jojo Sap-P 0.9 0.8 Cyphon confusus (Brown) ' Sap-P | 0.3 0.5 Cyphon obscurus (Guérin-Menéville) ! Cyphon ruficollis (Say) Cyphon variabilis (Thunberg) * Scraptiidae Sap-P | 1 | 2 | 0.4 0.3 | 1] 0.2 | Sap-P | o [ol o jo] o jo] o1 | 0.3 Anaspidinae Anaspis flavipennis Haldeman Anaspis nigrina Csiki Anaspis rufa Say Myceto | 88.3} 7 |125.8| 8 | 175.5] 4 | 141.3] 11 Scraptiinae Canifa pallipes (Melsheimer) 1 Canifa pusilla (Haldeman) 8 Scydmaenidae Euconnus debilitans (Casey) 0 Euconnus testaceipes (Casey) 2 Stenichmus badius (Casey) 4 Silvanidae Brontinae Dendrophagus cygnaei Mannerheim Myceto | 0 | O | 0.3 | 2 Or a}, 0 |r Onl (BEL. Silvaninae Silvanus bidentatus (Fabricius) ¢ Myceto 0 O°) “0.1 |. J 0 0 | 0.4 2 Sphinididae Odontosphindinae Odontosphindus denticollis LeConte | Myxo | 0 | 0; O 0 | 0 | 0 | 0.3 | 2 Sphinidinae Eurysphindus hirtus LeConte Myxo 0; 0.1 | 1 0 | 0 0 0 Sphindus trinifer Casey | Myxo | 0. r ia || OT, HO: | EGH Gale | uel Staphylinidae Aleocharinae Amischa analis (Gravenhorst) + Pred 0 0 0 0 O02 2 Atheta brunswickensis Klimaszewski aie Gin sc So oe Atheta dadopora Thomson + serra CEE acid elas Atheta hampshirensis (Bernhauer) Pred 0 0 0 0 0 Atheta klagesi Bernhauer * Pred Ge let) Ep WO sse5) se | PO ia) lO. Sa |EBES Atheta pennsylvanica Bernhauer Pred 0 0 0 0 |? 03 Atheta species 2 1 | “05. 2 5 Atheta vetricosa Bernhauer Pred 0 dese oy | ll 0 0 0 0 Deinopsis harringtoni Casey Pred Vans SO || a0) Ors | O3y | 0 0 336 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Disturbance Classes Clearcut | Clearcut + | Fire origin Wind 11 stands thin 4 stands | disturbance 8 stands 8 stands Av.S |S Gyrophaena species | Pred [| 0 |o}o1/i/ o fol/oi} i Leptusa brevicollis Casey Pred 10 Leptusa canonica Casey Pred Qin | 204 | oO) Leptusa carolinensis Pace Pred 0.4} 2] 11 0 0 3 0 Trophic Category oOlN Leptusa gatineauensis Klimaszewski Pred Leptusa opaca Casey Pred 0.4 0.6 Mocyta breviscula (Maklin) Myllaena arcana Casey Oxypoda nicriceps Casey Philhygra species | Philhygra species 2 Phloeopora species Omaliinae Acidota crenata (Fabricius) * Hapalaraea hamata (Fauvel) Phloeonomus laesicollis (Maklin) Paederninae Lithocharis thoracica (Casey) Palaminus hudsonicus Casey Scopaeus notangulus Casey Phloeocharinae Charyphus picipennis (LeConte) Pselaphinae Batrisodes lineaticollis (Aubé) Bibloplectus integer (LeConte) Bibloporus bicanalis (Casey) Euplectus duryi Casey Euplectus elongatus Brendel Reichenbachia spatulifer Casey Scaphidinae Baeocera congenera (Casey) Scaphidium quadriguttatum Say Scaphisoma lacustris Casey Staphylininae Atrecus americanus (Casey) Atrecus macrocephalus (Nordmann) Bisnius blandus (Say) Erichsonius patella (Horn) Gabrius microphthalmus (Horn) Gabrius picipennis (Maklin) Gyrohypnus cambelli Smetana Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia 337 Disturbance Classes Clearcut | Clearcut + | Fire origin Wind 11 stands thin 4 stands | disturbance 8 stands 8 stands S | Av.S | S Av. S Ontholestes cingulatus (Gravenhorst) | Pred | 0 | COT PT Philonthus caeruleipennis Mannerheim | Pred | 0 [ol] o [0] Trophic Category 0.2 Platydracus cupripennis (Melsimer) Pred ©: ,| EO; 1 Platydracus viridanus Horn Pred 0; O Quedius canadensis (Casey) Pred 0 | 0.1 Quedius capucinus (Gravenhorst) Pred 0 | 0.1 Quedius curtipennis Bernhauer + Pred Ol, | od Quedius peregrinus (Gravenhorst) Quedius plagiatus Mannerheim — — me |RoOte |e GDIWIOl/OJR INIT le ljoOIVNslolmn —_ Quedius rusticus Smetana Tympanophorus puncticollis (Erichson) j=) — me TRO te fe LR fl Rs TR | Ole Tachyporinae Bryophacis smetanai (Campbell) Pred Onl || Me 05 Bryoporus rufescens LeConte Pred O54?) 4 ||eelst Lordithon quaesitor (Horn) 0 Mycetoporus consors LeConte Sepedophilus cinctulus (Erichson) Sepidophilus crassus (Gravenhorst) Sepidophilus littoreus (Linnaeus) + Tachinus fumipennis (Say) Tachinus luridus Erichson Tachyporus dispar (Paykull) + Tachyporus nitidulus (Fabricius) + Stenotrachelidae Cephaloinae Cephaloon lepturoides Newman Cephaloon ungulare LeConte Tenebrionidae Alleculinae Capnochroa fuliginosa (Melsheimer) Hymenorus niger (Melsheimer) 0.3 lsomira quadristriata (Couper) , 7 Mycetochara analis (LeConte) Mycetochara bicolor (Couper) Bolitophaginae Bolitophagus corticola Say Bolito OF | |: |Leoe Pos Zona | 1 Tetratominae Tetratoma tesselata Melsheimer | Bolito | 0.1 | 1) 0.1 | 1 | 0 | 0 | 0.1 | 1 Throscidae Throscinae Aulonothroscus constrictor (Say) WVivicetor” |! Sol 5 | 95 ]| 28.9) 6" |NE co wlleida |e | PO Trixagus carinicollis (Schaeffer) Myceto | 0.4 | 3 0 OG. | 20.3) | || 6:4 2 Trogossitidae Peltinae _Thymalus marginicollis Chevrolat___|_Bolito | 0 | 0| 0 |o] 0 [oj] oi] 1 1 Species of the genus Cyp/on in the Maritime Provinces are presently being revised by Majka and Klausnitzer (in preparation). These species names may change in this forthcoming revision. 2 ‘This undescribed species of Paratenetus (previously included within Paratenetus fus- cus LeConte) will be described in a forthcoming publication by P. Bouchard and Y. Bousquet (in preparation). Notes: Trophic Categories: Bolito, Bolitophagous; Myceto, Mycetophagous; Myxo, Myxomycophagous; Pred, Predaceous; Rhizo, Rhizophagous; Sapro, Saprophytic; SE, Sap Feeder; Xylo, Xylophagous. f indicates a Palaearctic species; * indicates a Holarctic species. Deadwood and saproxylic beetle diversity in disturbed and managed spruce forests in Nova Scotia Appendix 2 Jo) SIMPER analysis showing average dissimilarity among sites by Disturbance Classes and the primary species of beetles involved. Clearcut and Clearcut and Thinned Average dissimilarity = 44.33 Clear-cut | CC @Thinned Species Av.Abund | Av.Abund Av.Diss | Diss/SD | Contrib% |Cum.% A. rufa 88.29 12575: 6.98 1.60 1325. 15375 I. quadristriata | 39.71 56.13 5.90 Lele 13.31 29.06 M. pumila 9.43 29.13 2.85 1.21 6.42 35.49 P ruficollis D7 i 28.63 2.39 0.67 5.30 40.78 Clearcut and Fire Disturbance Average dissimilarity = 54.51 Species Av.Abund | Av.Abund | Av.Diss | Diss/SD |Contrib% |Cum.% A. rufa 24.87 M. pumila 45.40 I. quadristriata |39.71 53752 Clearcut and thinned and Fire Disturbed Average dissimilarity = 58.09 Species Av.Abund Av.Abund | Av.Diss | Diss/SD | Contrib% | Cum.% A. rufa 25.25. 175.50 1355 2.08 23.33 23.33 M. pumila 25.13 89.50 10.49 E23 18.05 41.38 I. quadristriata 5.6.3 28.50 6.83 1.05 11375 po leo) Clear-cut and Wind Disturbed Average dissimilarity = 46.78 Species A, rufa Av.Abund Clearcut Wind Av.Abund Av. Diss Diss/SD Contrib% Cum.% I. quadristriata M. pumila P ruficollis 340 DeLancey J. Bishop et al. / ZooKeys 22: 309-340 (2009) Clear-cut and thinned and Wind Disturbed Average dissimilarity = 51.12 CC&Thinned | Wind | Species Av.Abund Av.Diss_ | Diss/SD | Contrib% | Cum.% A. rufa 125.75 913. |1.19 [17.86 17.86 I. quadristriata | 506.13 26.18 6.77 ia ke 13.24 BEN EO M. pumila 2513 12.00 Fale, 1.21 GAT. BT oF P ruficollis 28.63 7.09 2.77 0.76 5.43 42.70 A. flavipennis 18.00 135 3.24 45.94 Fire and Wind Disturbed Average dissimilarity = 57.64 Fire Species Av.Abund | Av.Abund | Av.Diss | Diss/SD | Contrib% |Cum.% A. rufa 175.50 1.53 25.08 25.08 M. pumila 89.50 1.24 18.54 43.63 I. quadristriata | 28.50 49.54 A. rufa = Anaspis rufa; I. quadristriata = Isomira quadristriata; M. pumila = Melanoph- thalma pumila; P ruficollis = Pidonia ruficollis; A. flavipennis = Anaspis flavipennis.