Supplementary material Supplement S1: Details on sampling design Standardized sampling protocol




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Supplementary material

Supplement S1: Details on sampling design

Standardized sampling protocol

The data used in the present study were complied from five different studies conducted in one ecoregion, each using a standardized sampling protocol. The beetle and true bug collecting effort was standardized for each trap. This sampling design was tested and standardized during the preparation of the first study in 1996 and then used in all following projects. This enabled us to make valid statistical comparisons of beetle and true bug diversity across scales, even though the samples contained different numbers of individuals (Summerville and Crist 2008). We are aware of a possible bias owing to sampling in different years. However, the samples collected with the same traps for 2 years in forest site HI had almost identical patterns of richness and community composition, and long-term studies from Finnish forest (Martikainen and Kouki, 2003) indicate that the patterns of beetle data collected in different stands by trapping during a whole vegetation period are consistent between sites even among different years (see Müller and Gossner 2010). Thus, this sampling design is more robust than short-time samplings, such as fogging campaigns or 1-month trappings, which are more sensitive to weather conditions and shifts in phenology, but are often used in beetle diversity studies (e.g., Gering et al., 2003; Grimbacher et al., 2006).



Description of sampled forest stands

The five studied ecoergions differ in their present-day forest composition, topography, macroclimate, and soil types. The forest site of the ecoregion Mainfränkische Platte (forest site Uffenheim; UF) is dominated by mixed oak forests. The forest sites of the ecoregion Fränkischer Keuper (forest sites Ebrach and Fabrickschlaichach; EB and FS) comprise two of the largest beech/oak-dominated forests in Germany. The forest site of the ecoregion Frankenalb (forest site Hienheim; HI) partly represents the primeval broadleaf tree species composition, but some spruce plantations are also included. In the ecoregion Tertiäres Hügelland (forest sites Krumbach and Ottobeuren; KB and OB), the favorable climate and soil conditions lead to a high productivity, resulting in a large-scale conversion of broadleaf to fast-growing, spruce-dominated forests, including also a high percentage of non-native Douglas fir (Gossner and Ammer 2006). The ecoregion Bayerischer Wald (forest site Bayerischer Wald; BW) shows an elevation gradient with mixed-montane forests at lower elevations and pure spruce stands at higher elevations. Management of these forests started historically rather late compared with other regions in Germany (Müller et al. 2008). The most natural forest landscapes, with respect to tree species composition, are in study sites UF, EB, FS, and BW. Details on the studied stands are given in Table S1.



Table S1: Description of the 28 mature forest stands (age >80 years) used for multiplicative and additive partitioning of true bug and saproxylic beetle diversity on five hierarchical levels. The ecoregion classification follows that of Walentowski et al. (2006), which is based on similar geological and climatic conditions for forest growth. The data is compiled from five different projects which followed a standardized sampling protocol (see paragraph “standardized sampling protocol”). SPFR = strictly protected forest research (from Müller and Gossner 2010)


Ecoregion/year of sampling

Forest site

Forest stand/altitude [m]

Number of traps

Stand type

Tree species (canopy traps)

Mainfränkische Platte / 2002


Uffenheim (UF)

A / 395

10

Open coppiced with standards, dominated by oak and hornbeam (height 18–25 m; age 140–170 years)

Quercus robur




B / 385

10

Coppiced with standards of oak, and a dense understory of aspen and hornbeam (height 18–24 m; age 140–170 years)

Quercus robur




C / 380

10

Closed conversion forest with oak, birch, and hornbeam (height 20–28 m; age 150–180 years)

Quercus robur




D / 380

10

Semi-open oak conversion forest with hornbeam and limb (height 21–29 m; 150–180 years)

Quercus robur

Fränkischer Keuper / 2004


Ebrach (EB)

A / 390

10

Beech dominated with interspersed oak trees (height: 25–38 m; age: 80–140 years)

Quercus petraea, Fagus sylvatica




B / 410

10

Beech dominated with interspersed oak and hornbeam trees (SPFR) (height: 28–43 m; age: 130–300 years)

Quercus petraea, Fagus sylvatica




C / 400

10

Beech dominated with interspersed oak and spruce trees (SPFR) (height: 28–43 m; age: 145–280 years)

Quercus petraea, Fagus sylvatica




D / 430

10

Beech dominated with interspersed oak trees (height: 26–38 m; age: 140–180 years)

Quercus petraea, Fagus sylvatica






Fabrickschlaichach (FS)

A / 420

10

Beech dominated with interspersed oak trees (height: 25–37 m; age: 100–140 years)

Quercus petraea, Fagus sylvatica




B / 410

10

Beech dominated with interspersed oak trees (height: 26–38 m; age: 140–175 years)

Quercus petraea, Fagus sylvatica




C / 400

10

Beech dominated with interspersed oak trees (height: 25–35 m; age: 80–130 years)

Quercus petraea, Fagus sylvatica




D / 410

10

Beech dominated with interspersed oak trees (height: 25–37 m; age: 140–180 years)

Quercus petraea, Fagus sylvatica

Frankenalb / 1997


Hienheim (HI)

A / 450

10

Spruce dominated with interspersed beech trees (height: 30–35 m; age 55–111 years)

Fagus sylvatica, Picea abies




B / 430

10

Spruce–beech–larch mixed forest (height: 28–36 m; age 91–111 years)

Fagus sylvatica, Picea abies, Larix decidua




C / 445

10

Beech–oak dominated with interspersed larch trees (height: 31–39 m; age 95–115 years)

Fagus sylvatica, Quercus petraea, Larix decidua




D / 420

10

Beech–oak dominated with interspersed spruce trees (SPFR) (height: 34–40 m; age 120–157 years)

Fagus sylvatica, Quercus petraea, Picea abies

Bayerischer Wald / 2007

Bayerischer Wald (BW)

A / 850

10

Mixed montane beech–silver fir–spruce forest (height 26–45 m; age 110–220 years)

Fagus sylvatica, Picea abies, Abies alba




B / 810

10

Mixed montane beech–silver fir–spruce forest (height 32–44 m; age 140–300 years)

Fagus sylvatica, Picea abies, Abies alba




C / 1250

10

High-montane spruce forests with interspersed mountain ash, maple (height 20–26 m; age 130–200 years)

Picea abies




D / 1250

10

High-montane spruce forests with interspersed mountain ash, maple (height 17–33 m; age 170–240 years)

Picea abies



Tertiäres Hügelland / 2000

Krumbach (KB)

A / 530

6

Spruce dominated with interspersed beech (height 36–39 m; age 70–100 years)

Fagus sylvatica, Picea abies




B / 530

6

Mixed spruce–beech with interspersed oaks (height 22–36 m; age 80–115 years)

Fagus sylvatica, Quercus robur, Picea abies




C / 520

6

Mixed beech–oak with interspersed spruce (height 26–35 m; age 85–148 years)

Fagus sylvatica, Quercus robur, Picea abies




D / 530

6

Oak dominated with interspersed beech and spruce (SPFR) (height 23–37 m; age 110–170 years)

Fagus sylvatica, Quercus robur, Picea abies

Ottobeuren (OB)

A / 630

6

Spruce dominated with interspersed beech and Douglas fir (height 29–43 m; age 80–105 years)

Fagus sylvatica, Picea abies, Pseudotsuga menziesii




B / 625

6

Mixed spruce-beech with interspersed Douglas fir (height 28–45 m; age 80–116 years)

Fagus sylvatica, Picea abies, Pseudotsuga menziesii




C / 645

6

Beech dominated with interspersed spruce and Douglas fir (Height 35–45 m; age 101–134 years)

Fagus sylvatica, Picea abies, Pseudotsuga menziesii




D / 640

6

Beech dominated with interspersed spruce and Douglas fir (SPFR) (height: 34–45 m; age 108–118 years)

Fagus sylvatica, Picea abies, Pseudotsuga menziesii

Figure S1: Location of the 28 stands studied at seven forest sites within five Bavarian ecoregions. Inset: location of Bavaria in Germany. Forest sites: Uffenheim (UF), Fabrickschlaichach (FS), Ebrach (EB), Hienheim (HI), Krumbach (KB), Ottobeuren (OB), Bayerischer Wald (BW). Gray shading indicates forested area. Lines indicate borders between forest ecoregions

Supplement S2: Species identification

All species were identified by viewing with a stereomicroscope. True bugs species were identified by the first author (MMG), except for specimens from site Hienheim (identified by T. Maier, München), using common identification keys (Wagner 1952, 1966, 1967; Péricart 1972, 1983, 1987, 1990, 1998) and several other species-level publications. All saproxylic beetles were identified to the species level by external taxonomic specialists (H. Bussler, Feuchtwangen; F. Köhler, Bornheim; B. Büche, Berlin) according to Freude et al. (1964–1983) and several other species-level publications.



Supplement S3: Body-size distributions



Figure S3: Body-size distribution of all (a) true bug and (c) saproxylic species recorded in Germany and of all sampled (b) true bug and (d) saproxylic species sampled. Each species is classified into one of three body-size classes with equal number of species. Class borders are indicated by dashed lines. In the present study, small, medium, and large species were classified based on the distribution of all species recorded in Germany

Supplement S4: Additive and multiplicative diversity partitioning

Additive partitioning revealed a much higher proportion of the highest β-diversity level and a lower α-diversity proportion compared to the multiplicative approach (see Figs. S4-1 and S4-2). Moreover, the intermediate spatial scale (especially stand level) was much more important in the additive approach, while small spatial scales (especially trap level) were of higher significance in the multiplicative approach. The main patterns among functional guilds were similar, except for the host specificity of saproxylic beetles, for which the α-diversity was most important for medium-sized species in the multiplicative approach, but not in the additive approach.

We tested the statistical significance of level-specific α- and β-diversity estimates using a randomization procedure. We used an unrestricted individual-based randomization (Crist et al. 2003; Veech and Crist 2009) to generate 10,000 random distributions of true bug and saproxylic beetle species by reassigning each individual of the dataset to any other sample. This results in a null distribution of each α- and β-diversity estimate for each level. Each of the original level-specific estimates was then compared with the appropriate null distribution and used to test the null hypothesis that the observed α- and β-diversities are obtained by a random distribution of individuals among samples at all hierarchical levels. Statistical significance was assessed by the proportion of null values that are greater than (or less than) the actual estimate (Manly 1997; Roff 2006). In both additive and multiplicative partitioning, a significantly higher proportion of β-diversity of the highest level, among-ecoregions, and a lower proportion of α-diversity than expected by chance were observed (see Table S4-1).

For both true bugs and saproxylic beetles, a higher proportion of α-diversity and a lower proportion of β-diversity than expected by chance were found. This indicates a patchy distribution of true bugs and saproxylic beetles in Central European forests, reflecting intraspecific aggregation among patches as predicted by He and Legendre (2002). In contrast to the results of Summerville et al. (2006a) on forest Lepidoptera in deciduous forests of North America, a higher β-diversity than expected by chance was observed only on the highest scale (ecoregion level) when a comparable additive approach was applied. This suggests that there is also a species aggregation among strata, stands, and forests. This, however, might be due to the methodological limitations of the additive approach (see Jost 2007). When a multiplicative approach was applied, a higher β-diversity than expected by chance was observed on these intermediate spatial scales. This was confirmed for both taxa and all functional guilds (differences in body size or host specificity). This leads to the conclusion that intraspecific aggregation generally occurs mainly on very small spatial scales.



Table S4-1: Significance values for tests of diversity estimates from additive or multiplicative partitioning against null estimates using unrestricted individual-based randomizations. Abbreviations: - significantly small, + significantly large, ns not significant at the 0.05 level








True bugs




Saproxylic beetles







Additive

Multiplicative




Additive

Multiplicative

Group

Level

Species

richness

q =0

q = 0.999

q = 2




Species

richness

q = 0

q = 0.999

q = 2

Body size































Small

β 5

<0.001 (+)

<0.001 (+)

<0.001 (+)

<0.001 (+)




<0.001
(+)

<0.001 (+)

<0.001 (+)

<0.001 (+)




β 4

ns

<0.001
(-)

0.001

(+)


ns




<0.001
(-)

0.006
(-)

0.003
(-)

ns




β 3

<0.001
(-)

<0.001
(-)

ns

ns




<0.001
(-)

<0.001 (+)

<0.001
(-)

<0.001 (-)




β 2

<0.001
(-)

ns

<0.001
(-)

<0.001 (-)




<0.001
(-)

<0.001 (+)

<0.001 (+)

<0.001 (+)




β 1

<0.001
(-)

<0.001
(-)

<0.001
(-)

0.005 (+)




<0.001
(-)

<0.001 (+)

<0.001
(-)

<0.001 (-)




α 1

<0.001
(-)

<0.001
(-)

<0.001
(-)

<0.001 (-)




<0.001
(-)

<0.001
(-)

<0.001
(-)

<0.001 (-)

Medium

β 5

<0.001 (+)

<0.001 (+)

<0.001 (+)

<0.001 (+)




<0.001
(+)

<0.001 (+)

<0.001 (+)

<0.001 (+)




β 4

ns

<0.001 (+)

0.008
(+)

ns




<0.001
(-)

0.003
(+)

0.003
(+)

0.001 (+)




β 3

0.002
(-)

<0.001 (+)

ns

ns




<0.001
(-)

<0.001 (+)

<0.001
(-)

<0.001 (+)




β 2

0.002
(-)

<0.001 (+)

ns

ns




<0.001
(-)

<0.001 (+)

<0.001 (+)

ns




β 1

<0.001
(-)

<0.001 (+)

0.001

(-)


<0.001 (-)




<0.001
(-)

ns

<0.001
(-)

<0.001 (+)




α 1

<0.001
(-)

<0.001
(-)

<0.001
(-)

<0.001 (-)




<0.001
(-)

<0.001
(-)

<0.001
(-)

<0.001 (-)

Large

β 5

<0.001 (+)

<0.001 (+)

<0.001 (+)

<0.001 (+)




<0.001
(+)

<0.001 (+)

<0.001 (+)

<0.001 (+)




β 4

<0.001
(-)

<0.001
(-)

ns

ns




0.001
(-)

0.015
(+)

<0.001 (+)

ns




β 3

<0.001
(-)

0.002
(+)

ns

ns




<0.001
(-)

0.002
(+)

0.005
(-)

ns




β 2

<0.001
(-)

<0.001 (+)

ns

ns




<0.001
(-)

<0.001 (+)

<0.001
(-)

<0.001 (-)




β 1

<0.001
(-)

ns

<0.001
(-)

<0.001 (-)




<0.001
(-)

<0.001
(-)

<0.001
(-)

<0.001 (-)




α 1

<0.001
(-)

<0.001
(-)

<0.001
(-)

<0.001 (-)




<0.001
(-)

<0.001
(-)

<0.001
(-)

<0.001 (-)
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