Biological Conservation 143 (2010) 625–633

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Biological Conservation
journal homepage: www.elsevier.com/locate/biocon

Three-dimensional partitioning of diversity informs state-wide strategies

for the conservation of saproxylic beetles
Jörg Müller a,*, Martin M. Goßner b,1
a
Bavarian Forest National Park, Freyunger Str. 2, D-94481 Grafenau, Germany
b
Institute of Ecology, Friedrich-Schiller-University, Dornburger Str. 159, D-07743 Jena, Germany

a r t i c l e i n f o a b s t r a c t

Article history: Global conventions on biological diversity force governments to develop region-wide conservation strat-
Received 3 June 2009 egies. Such strategies are difficult to design for all taxa because little is known about the important spatial
Received in revised form 23 November 2009 scales. Here we applied additive partitioning of the diversity of saproxylic beetles in Bavarian forests in
Accepted 25 November 2009
Southern Germany using a nested hierarchical design of five increasingly broader spatial levels: trap,
Available online 23 December 2009
strata, forest stand, forest site, and ecoregion. We consistently found a significantly higher percentage
than expected by chance of between ecoregion diversity and significantly lower a diversity within traps.
Keywords:
A significant proportion of b diversity was also found between stands. Analysis of species represented by
Canopy diversity
Critical diversity scales
<0.005% of all specimens in our samples and of species classified as threatened revealed similar results.
Natura 2000 Critical spatial scales for threatened species encompassed the critical levels of common species. Within
Null hypothesis habitat substratum guilds, the proportion of b diversity increased from species associated with fresh
Species richness wood to those associated with rotten wood to those associated with fungi. Our results suggest that the
Strictly protected areas most effective way to ensure saproxylic beetle diversity in a state-wide strategy is to add new conserva-
tion sites within different ecoregions and to establish new conservation areas in additional forest stands,
rather than to enlarge reserves. Our findings further suggest that monitoring of saproxylic beetle diver-
sity on a broader scale in European temperate forests can be restricted to ‘‘monitoring species”, i.e. a sub-
group of families easy to identify, and that canopy sampling can be neglected without a substantial loss of
information.
Ó 2009 Elsevier Ltd. All rights reserved.

1. Introduction
Since our understanding that hyperdiverse taxa such as arthro-
pods fulfil important ecosystem functions and services, conserva-
tion biology changed its focus from single species to ecosystem
management (Lister, 1998; Weisser and Siemann, 2004). Global
discussions on the loss of diversity have forced governmental
administrations to rapidly develop biodiversity policies (e.g. the
network European Natura 2000), unfortunately often without suf-
ficient knowledge of the organization of biodiversity on spatial
scales (Chiarucci et al., 2008; Haslett, 2007). The recently-pub-
lished European Strategy for the Conservation of Invertebrates
compiles substantial information on the conservation needs of
the world’s most-documented insect fauna, with the goal to stop
their decline; seven main objectives to enhance invertebrate con-
servation in Europe are listed (Haslett, 2007). Two of these objec-
tives (‘‘2. Promote integrated management of landscape mosaics
* Corresponding author. Tel.: +49 8552 9600 179; fax: +49 8552 9600100.
E-mail addresses: joerg.mueller@npv-bw.bayern.de (J. Müller), martin.gossner@
uni-jena.de (M.M. Goßner).
1
Tel.: +49 3641 949481; fax: +49 3641 949402.
at the relevant scales. . .” and ‘‘4. Identify and prioritise key actions
to be implemented at different political and geographical levels”)
entail the identification of relevant scales for such strategies.
Forests in Central Europe, once the major primeval habitat, have
greatly decreased because of cultivation, resulting in a mosaic of
forest patches of different size, connectivity and distribution
(Rüther and Walentowski, 2008). The distribution of species
assemblages in space can be understood by focusing on the
ecological processes of species interactions and dispersal within
the framework of metacommunity ecology (Hanski, 2004; Leibold,
2004). Dispersal limits have been enforced by human fragmenta-
tion of the landscape, which results in forests mostly surrounded
by highly contrasting environmental matrices (Ricklefs, 1987;
Shepherd and Brantley, 2005). The hyperdiverse saproxylic beetles
are particularly suitable for such studies for a number of reasons:
they are species rich; they are sensitive to logging; they have a
broad functional importance in forest ecosystems, i.e. as fungi vec-
tors, predators, xylophages, and detritivores (Davies et al., 2008;
Grove, 2002; Speight, 1989); they are under focus in conservation,
resulting in an overrepresentation in many red lists (Brustel, 2004;
Schmidl et al., 2003); and the taxonomical literature allows almost
all specimens collected in the field to be identified to the species

0006-3207/$ - see front matter Ó 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2009.11.027
626 J. Müller, M.M. Goßner / Biological Conservation 143 (2010) 625–633
level (Freude et al., 1964–1983). Particular families of saproxylic
beetles have been classified as suitable for monitoring of conserva-
tion status in landscape ecological studies of Central Europe be-
cause they are relatively easy to determine and ecologically
representative (Schmidl and Bußler, 2004). Here, we refer to these
beetles as ‘‘monitoring species”.
Despite this relatively high state of knowledge, little is known
about the biodiversity patterns at various spatial scales because
most studies have been restricted to a few stands or sites (c.f.
Davies et al., 2008). Even studies that proposed theoretical models
of long-term suitability of saproxylic organisms have been
restricted to relatively small, local areas (Tikkanen et al., 2007).
Understanding the influence of spatial scales on diversity patterns,
however, is absolutely essential for developing a state-wide con-
servation strategy and selecting additional sites for conservation,
to focus on an effective effort of local forest managers and an effec-
tive monitoring of biodiversity (Chiarucci et al., 2008). Important
levels of forest arthropod diversity range in scale from microsites
to stands and from near ground to canopy, to forests, among for-
ests, and among ecoregions; these levels have been discussed but
never analysed in one multi-scaled study.
Here we investigated which spatial level is critical for the total
diversity of saproxylic beetles. We compiled standardized data on
saproxylic beetles sampled in the state of Bavaria in Germany. The
areas sampled included several ecoregions and different types of
forests, from forests dominated by lowland oak to those dominated
by mountain spruce. We tested the hypothesis that broad-scale dif-
ferences between forests and ecoregions are more important in
influencing saproxylic beetle diversity and community composi-
tion than local differences among trees and stands within forests
and ecoregions. Our null hypothesis was that the observed diver-
sity across hierarchical levels does not differ from that expected
by random distribution of individuals. Our alternative hypotheses
predicted significant deviance of diversity estimates from random
expectation at: (1) fine spatial scales owing to differences in spe-
cies composition and abundance among traps within stands and
among strata, (2) intermediate spatial scales owing to differences
among stands within a forest, and (3) broad spatial scales owing
to differences between forests and between ecoregions. We also
examined the contrasting roles of: (1) common species and those
considered to be threatened according to the state-wide red list
in comparison to abundant and rare species in our samples, (2)
‘‘monitoring species”, a subgroup relatively easy to determine
and ecologically representative, and (3) three habitat substratum
guilds in their contribution to the scaling of species diversity and
composition. Our goal was to identify how differences in subsets
of species affect the critical levels for the diversity of saproxylic
beetle community structure. Based on our results, we provide sug-
gestions for the spatial establishment of new strictly protected for-
est reserves state wide, the management of biodiversity on the
local scale, and biodiversity assessment and monitoring for the
state.
2. Methods
2.1. Study sites and sampling design
Beetles were sampled following a standardized protocol
(Summerville and Crist, 2008) from 1996 to 2007 in different pro-
jects. The data were compiled to create a hierarchically nested ma-
trix that included the main ecological forest types of Southern
Germany, except the floodplain forests. The five hierarchical levels
corresponded to spatial scales: ecoregion, forest site (connected
forest area >500 ha), stand (5–70 ha), and two strata within the
stand (near ground and canopy). The highest level and therefore
the broadest spatial scale was represented by five ecoregions
(Walentowski et al., 2004) (Table 1 and Fig. 1) that 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 repre-
sents 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 favourable climate and soil conditions lead to a high productiv-
ity, resulting in a large-scale conversion of broadleaf to fast-grow-
ing, 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 ele-
vation 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 land-
scapes, with respect to tree species composition, are in study sites
UF, EB, FS, and BW.
2.2. Beetle data
Saproxylic beetle communities were sampled over one whole
vegetation period, i.e. from April to October, in most areas, except
in high montane stands, where sampling started immediately after
the snow melt in May. Hence, each sample represents the commu-
nity of a whole year. We used flight-interception traps consisting of
a crossed pair of transparent plastic shields (40  60 cm), as de-
scribed by Hyvärinen et al. (2006).
In each stand, five trees representing the dominating tree spe-
cies in the overstory were randomly selected (Table 1). Traps were
installed pair-wise in the selected tree crowns and near the
ground. In the forest sites OB and KB, only three pairs of traps were
installed in each stand. Canopy traps were positioned in the centre
of the tree crowns in the upper canopy layer either by using a
crossbow or by climbing the tree; near-ground traps were hung
on a rope 1.5 m above ground between two trees. Collected beetles
were killed and preserved in a non-attracting 3% copper vitriol
solution. All saproxylic beetles were identified to the species level
according to Freude et al. (1964–1983). Saproxylic beetles are
defined as those beetles that depend on dead or dying wood of
moribund or dead trees, on wood-inhabiting fungi, or on the pres-
ence of other saproxylic organisms during some part of their life
cycle (Speight, 1989).
The beetle collecting effort was standardized for each trap. This
sampling design was tested and standardized during the prepara-
tion of the first study in 1996 and then used in all following pro-
jects. This enabled us to make valid statistical comparisons of
beetle 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 beetle data col-
lected in different stands by trapping during a whole vegetation
period reveals consistent patterns between sites even among dif-
ferent years (see Supplementary material). Thus, this sampling de-
sign 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
 J. Müller, M.M. Goßner / Biological Conservation 143 (2010) 625–633 627

Table 1
Description of the 28 forest stands used for additive partitioning of saproxylic beetle diversity on five hierarchical levels. SPFR = strictly protected forest research.

Ecoregion/year of Forest site Forest stand/ Number of Stand type Tree species (canopy traps)
sampling altitude (m) traps
Mainfränkische Uffenheim (UF) A/395 10 Open coppiced with standards, dominated by oak and Quercus robur
Platte/2002 hornbeam
B/385 10 Coppiced with standards of oak, and a dense Quercus robur
understory of aspen and hornbeam
C/380 10 Closed conversion forest with oak, birch and Quercus robur
hornbeam
D/380 10 Semi-open oak conversion forest with hornbeam and Quercus robur
limb
Fränkischer Ebrach (EB) A/390 10 Beech dominated with interspersed oak trees Quercus petraea, Fagus sylvatica
Keuper/2004
B/410 10 Beech dominated with interspersed oak and Quercus petraea, Fagus sylvatica
hornbeam trees (SPFR)
C/400 10 Beech dominated with interspersed oak and spruce Quercus petraea, Fagus sylvatica
trees (SPFR)
D/430 10 Beech dominated with interspersed oak trees Quercus petraea, Fagus sylvatica
Fabrickschlaichach A/420 10 Beech dominated with interspersed oak trees Quercus petraea, Fagus sylvatica
(FS)
B/410 10 Beech dominated with interspersed oak trees Quercus petraea, Fagus sylvatica
C/400 10 Beech dominated with interspersed oak trees Quercus petraea, Fagus sylvatica
D/ 410 10 Beech dominated with interspersed oak trees Quercus petraea, Fagus sylvatica
Frankenalb/1997 Hienheim (HI) A/450 10 Spruce dominated with interspersed beech trees Fagus sylvatica, Picea abies
B/430 10 Spruce-beech-larch mixed forest Fagus sylvatica, Picea abies, Larix
decidua
C/445 10 Beech-oak dominated with interspersed larch trees Fagus sylvatica, Quercus petraea,
Larix decidua
D/420 10 Beech-oak dominated with interspersed spruce trees Fagus sylvatica, Quercus petraea,
(SPFR) Picea abies
Bayerischer Wald/ Bayerischer Wald A/850 10 Mixed montane beech-silver fir-spruce forest Fagus sylvatica, Picea abies, Abies
2007 (BW) alba
B/810 10 Mixed montane beech-silver fir-spruce forest Fagus sylvatica, Picea abies, Abies
alba
C/1250 10 High-montane spruce forests with interspersed Picea abies
mountain ash, maple
D/1250 10 High-montane spruce forests with interspersed Picea abies
mountain ash, maple
Tertiäres Krumbach (KB) A/530 6 Spruce dominated with interspersed beech Fagus sylvatica, Picea abies
Hügelland/
2000
B/530 6 Mixed spruce-beech with interspersed oaks Fagus sylvatica, Quercus robur,
Picea abies
C/520 6 Mixed beech-oak with interspersed spruce Fagus sylvatica, Quercus robur,
Picea abies
D/530 6 Oak dominated with interspersed beech and spruce Fagus sylvatica, Quercus robur,
(SPFR) Picea abies
Ottobeuren (OB) A/630 6 Spruce dominated with interspersed beech and Fagus sylvatica, Picea abies,
Douglas fir Pseudotsuga 0menziesii
B/625 6 Mixed spruce-beech with interspersed Douglas fir Fagus sylvatica, Picea abies,
Pseudotsuga menziesii
C/645 6 Beech dominated with interspersed spruce and Fagus sylvatica, Picea abies,
Douglas fir Pseudotsuga menziesii
D/640 6 Beech dominated with interspersed spruce and Fagus sylvatica, Picea abies,
Douglas fir (SPFR) Pseudotsuga menziesii

beetle diversity studies (Bußler et al., 2004; Gering et al., 2003;
Grimbacher et al., 2006).
In addition to analysing the entire community, we also selected
several subsets of the community to assess their relationship to
different scales of diversity. First, we classified species as being
common or threatened according to the IUCN classifications of
the recent red list of Bavaria; this list includes all beetle families
whose members are classified as saproxylic (Schmidl et al.,
2003). Second, to examine whether similar results were obtained
with rare and common species, we classified species as being of
high density, i.e. consisting of >0.5% of the whole density, and as
being of low density, i.e. consisting of <0.005% of the whole den-
sity, according to Gering et al. (2003). Third, diversity patterns of
‘‘monitoring species” suggested for state-wide monitoring accord-
ing to Schmidl et al. (2003) were assessed in comparison to the
diversity patterns of the entire community. Finally, we separated
the data set according to the three major habitat substratum guilds
of saproxylic species: fresh dead wood, extensively rotten dead
wood, and wood-inhabiting fungi (Schmidl and Bußler, 2004).
Two of the substratum guilds proposed by Schmidl and Bußler
(2004), ‘‘living in rotholes” and ‘‘others”, were excluded in the
analysis because of the low number of species sampled within
these guilds.
2.3. Data analysis
Patterns in beetle species composition among the 28 forest
stands were displayed with non-metric multidimensional scaling
ordination (nMDS) using the software vegan in R 2.8.0 (R Develop-
ment Team 2008). We constructed species-accumulation curves
628 J. Müller, M.M. Goßner / Biological Conservation 143 (2010) 625–633

within the vegan package (Oksanen et al., 2006). Using this func-
tion, mean values of a diversity at lower levels of a sampling hier-
archy are compared to the total number of species in the entire
data set (c diversity) in the form: c = mean(a)+b. Thus, b = c–
mean(a). This additive procedure is extended across the a and five
b scales in a hierarchical design described above (for more infor-
mation about the hierarchical designs in diversity partitioning,
see Gering et al., 2003 and Fig. S1). Average a components were
weighted with sample abundances to calculate the weighted
averages.
The statistical significance of level-specific a and b estimates
can be tested within the adipart function through a randomization
procedure. We used a complete randomization (Crist et al., 2003)
to generate 10,000 random distributions of saproxylic beetle spe-
cies among samples at all hierarchical levels to generate the null
distribution of each a and b estimate for each level. Each of the ori-
ginal level-specific estimates was then compared with the appro-
priate null distribution and used to test the null hypothesis that
the observed a and b diversity are obtained by a random distribu-
tion of individuals among samples at all hierarchical levels. Statis-
tical significance was assessed by the proportion of null values that
are greater than (or smaller than) the actual estimate (Manly,
1997; Roff, 2006).

3. Results
Fig. 1. Location of the seven forest sites within the five Bavarian ecoregions studied.
Inset: location of Bavaria in Germany. Forest sites: Uffenheim (UF), Fabrickschla- 3.1. General patterns
ichach (FS), Ebrach (EB), Hienheim (HI), Krumbach (KB), Ottobeuren (OB), Bayeri-
scher Wald (BW). Grey shading indicates forested area. Lines indicate borders
Our final data set comprised 23,985 individuals representing
between forest ecoregions.
470 species of saproxylic beetles. Two of these species (0.4%) were
associated with one tree species, and 61 species (13%) were associ-
using the function specaccum in the vegan package with the meth- ated with one tree genus. However, the majority of the species
od rarefaction. To test for differences in pairs of traps (near ground were generally associated with broad-leaved trees or conifers or
and canopy), we used a pair-wise Wilcoxon test. with both (see Supplementary material Table S1). The non-metric
Whittaker (1960, 1977) showed that diversity can be separated multidimensional scaling showed a clear separation of the differ-
in a (within sampling units), b (among units), and c (regional) ent ecoregions along the first dimension from xerothermic oak
diversity. Allan (1975) and Lande (1996) demonstrated that regio- forests to beech-dominated forests at lower elevations to spruce-
nal diversity is the sum of a and b diversity when a is the average dominated forests (Fig. 2; left to right). The forests dominated by
diversity within the sampling units in the region and b is the diver- spruce plantations differed from montane forests along the second
sity among sampling units. This additive procedure can be freely dimension. The stands within an ecoregion were more clustered
extended across multiple scales in a hierarchical sampling design than stands between ecoregions. This pattern was highly stable
with i = 1, 2, 3, . . . , m levels of sampling (Crist et al., 2003). This ap-
proach allows the additive partition of the total diversity in a re-
1.0
gion into scale-specific diversity components, which can be UF
D
directly compared. In the context of our study, the overall beetle A
EB
diversity can be described by the following formula: B
C
FS
0.5 B HI
C A
B
KB
c ¼ a1 ðwithin trapÞ þ b1 ðamong trapsÞ þ b2 ðamong strataÞ D D A OB
C BW
NMDS2

þ b3 ðamong standsÞ þ b4 ðamong forestsÞ

0.0 A
D
þ b5 ðamong ecoregionsÞ:
C A
C D
B B B
Lande (1996) demonstrated that any metric can be partitioned D
A
D
into its components provided that it exhibits strict concavity, -0.5 C
C
which means that the overall value of that metric for a pooled B
A
set of communities equals or exceeds the average diversity within
communities. Species richness, Simpson diversity, and Shannon -1.0
diversity fulfil these criteria and were established in numerous -1.0 -0.5 0.0 0.5 1.0
additive partitioning studies. We used all three metrics in our NMDS1
study to account for the pure effects of species richness and the
combined effects of richness and abundance, with the Shannon in- Fig. 2. Non-metric multidimensional scaling ordination (stress value = 13.64) of
dex being more sensitive and the Simpson diversity being less sen- beetle species composition using presence/absence data (470 species; 23,985
individuals) of 28 forest stands within seven forests (UF, Uffenheim; EB, Ebrach; FS,
sitive to rare species in the samples (Gering et al., 2003; Lande, Fabrickschlaichach; HI, Hienheim; KB, Krumbach; OB, Ottobeuren; BW, Bayerischer
1996; Magurran, 2004). Using these diversity indices, we addi- Wald) in Bavaria. Symbols represent the forest site, letters refer to stands explained
tively partitioned the entire community using the adipart function in Table 1, and stands of one ecoregion are encircled.
 J. Müller, M.M. Goßner / Biological Conservation 143 (2010) 625–633 629

and independent of the use of presence/absence or square-root- 3.2. Additive partitioning

transformed abundance data (not shown).
Species richness was not distributed evenly among the ecore-
gions. The rarefaction curves (Fig. 3) showed a decreasing species
richness from oak-dominated forest landscapes (Mainfränkische
Platte) to forest landscapes dominated by natural tree species,
independent of the elevation (Fränkischer Keuper, Frankenalb,
Bayerischer Wald), to forests intensively used for commercial
wood production dominated by spruce plantations with non-na-
tive Douglas fir (Tertiäres Hügelland) (Fig. 3). A comparison of spe-
cies richness in canopy and near-ground traps showed a
significantly lower species richness in canopy traps. This was also
supported by a pair-wise Wilcoxon test (Fig. 4). The majority of
species with a preference for one of the two strata were associated
with the near-ground stratum (see Supplementary material Table
S2).
Additive partitioning revealed a constant and significantly high-
er b diversity proportion of the highest b level between ecoregions
than expected by chance, whereas the a component was always
lower than expected (Table 2, Fig. 5). This was evident across all
combinations of groups and diversity measures (Table 2), despite
the a component accounting for a substantial proportion of the
Table 2
Significance values for tests of diversity estimates from additive partitioning against
null estimates using complete randomization, obtained using the function adipart
within the vegan package, R2.8.1 system). All values were determined at the 0.005
level after Bonferoni correction. Abbreviations:  significantly small, + significantly
large, ns not significant.

Group Level Species Simpson Shannon

richness diversity diversity
Entire community b5 + + +
b4  + +
b3  + +
b2  + +
b1  + +
a1   
Red-listed species b5 + + +
b4  + +
b3  + +
b2  + +
b1  + 
a1   
Common species b5 + + +
b4 ns + +
b3  + +
b2 ns + +
b1  + 
a1   
Species with low density b5 + + +
(<0.005%) b4  + ns
b3  + 
Fig. 3. Accumulation curves for beetle species richness from 248 flight-interception b2  + ns
trap samples of five ecoregions, including seven forest sites, each containing four b1  + 
stands with five traps in the canopy and five traps near ground. Note that in a1   
Tertiäres Hügelland, the number of traps was only six per stand at both forest sites. Species with high density b5 + + +
Curves and 95% confidence intervals were calculated using the specaccum function (>0.5%) b4 + + +
in vegan, with the method rarefaction. The open circles indicate the species richness b3 + + +
at 2000 individuals. The inset shows the accumulation curve of all 248 traps. b2 + + +
b1 + + +
a1   
Monitoring species b5 + + +
b4 ns + +
b3  + +
b2  + +
b1  + 
a1   
Species associated with fresh b5 + + +
wood b4  + +
b3 ns + +
b2 + + +
b1  + 
a1   
Species associated with b5 + + +
advanced decomposed b4  + +
wood b3  + +
b2  + +
b1  + 
a1   
Species associated with wood- b5 + + +
inhabiting fungi b4 ns + +
Fig. 4. Rarefaction curves for beetle species richness from 248 traps (124 in the b3  + +
canopy and 124 near ground). Confidence intervals were calculated using the b2  + +
function specaccum with the method rarefaction within the vegan package (R2.8.1 b1  + ns
system). A pair-wise Wilcoxon test showed a significantly lower number of species a1   
in the canopy (V = 1628.5, p < 0.001, two-sided).
630 J. Müller, M.M. Goßner / Biological Conservation 143 (2010) 625–633

richness of abundant species (>0.5% of all specimens) in our sam-
ples (Fig. 5) and for a large proportion of the Shannon and Simpson
diversity for the entire community (Fig. 6). However, for threa-
tened and common species, the difference in observed proportions
of b diversity had the same direction as abundant and rare species,
and was higher among ecoregions than expected by chance, but
the difference between threatened and common species was much
lower. The ‘‘monitoring species” showed a highly similar pattern
compared to the entire community (Fig. 5). Among the substratum
guilds, the proportion of a diversity was highest for fresh-wood
species, while the proportion of diversity among ecoregions was
greatest for fungi-associated species. Two estimates of b diversity —
the diversity among strata and among forest sites — and also the a
diversity were consistently relatively low (Fig. 5). We also found
differences between Shannon and Simpson diversity in the per-
centage of total diversity that accounted for a diversity within
traps; the difference was greater for Simpson diversity than for
Shannon diversity (Fig. 6). Deviations from expected values were
positive for all components of b diversity and negative for all a
diversity.
et al., 2008; Summerville et al., 2003). Direly needed answers to
questions pertaining to establishing, enlarging, and monitoring of
broad-scale networks of protected areas, such as Natura 2000 in
Europe, can be developed using this approach (cf. Chiarucci et al.,
2008; Juranski et al., 2009).
4.1. Important spatial levels of biodiversity
How c diversity is partitioned into a and b diversity compo-
nents depends on the spatial scale of the observations (Lindo and
Winchester, 2008; Loreau, 2000) and has been linked to particular
ecological traits, such as dispersal ability, body size, or niche
breadth (Summerville et al., 2006). In our study, estimates of two
levels of diversity showed the same deviation from random and

4. Discussion

Although the habitat requirements and conservation status of

saproxylic beetles in Germany are very well known, our study is
the first to analyse how the diversity of saproxylic beetles of Euro-
pean temperate forests is partitioned on a state-wide scale. With
470 species of saproxylic beetles, our data set comprised 42% of
all species recorded from the state Bavaria (c.f. Köhler and Klausn-
itzer, 1998). Even if the overall species accumulation curve (Fig. 3
inset) did not show a saturation, it was flatter than those curves of
comparable beetle studies (c.f. Gering et al., 2003), even though we
included only 5 of 14 ecoregions in Bavaria in our analysis (see Fig. 6. Percentage of total Shannon and Simpson diversity explained by a and b
components of diversity for the whole community (470 species) on five spatial
Fig. 1). The high flexibility makes additive partitioning highly inter-
scales: traps, strata, forest stands, forest sites, and ecoregions. Contributions to total
esting for a regional (in the sense of a state) but multi-scaled con- diversity for each scale were determined by additive partitioning of diversity using
servation analysis (Gering et al., 2003; Hirao et al., 2007; Ribeiro the adipart function within the vegan package (R2.8.1 system).

Fig. 5. Percentage of total beetle species richness explained by a and b components of diversity on five spatial scales: traps, strata, forest stands, forests sites, and ecoregions.
The contributions to the total richness for each scale were determined by additive partitioning of diversity using the function adipart within the vegan package (R2.8.1
system). The total species richness for each of the groups denoted on the x-axis is indicated by the number at the top of each bar. Red-listed species were classified according
to the red list of Bavaria (Schmidl et al., 2003), which comprises all beetle families. Species not listed in the red list of Bavaria were classified as common species a priori.
Species with a low or a high density in our samples were classified according to Gering et al. (2003). ‘‘Monitoring species” were selected using the classification of species
relevant in landscape ecology planning (Schmidl and Bußler, 2004). Species colonizing three different dead-wood habitats were classified according to Schmidl and Bußler
(2004).
 J. Müller, M.M. Goßner / Biological Conservation 143 (2010) 625–633
were independent from the metric and the component of the com-
munity analysed: The a diversity within traps was always smaller
than expected by random chance, whereas the b diversity between
ecoregions was always higher than expected. The importance of
the ecoregion for overall diversity was also supported by our ordi-
nation, which identified strong differences in community composi-
tion among ecoregions (Fig. 3). Our results support general findings
that insect community composition varies greatly on larger scales,
while species dominance (Simpson diversity) in temperate forests
is determined on finer scales (Summerville et al., 2003). This is not
surprising because species have spatially limited and non-random
distributions (Morlon et al., 2008). b diversity in the sense of the
difference between the species compositions is thus strongly cor-
related with distance, but with different underlying reasons (Har-
rison et al., 1992; Rosenzweig, 1995). This suggests an increasing
b diversity with increasing spatial scale. The low contribution of
diversity among forests in our study, however, did not support this
expectation in its entirety and indicated that the correlation be-
tween species composition and distance is not linear.
The high importance of b diversity between ecoregions in our
study could be explained by three different factors. First, dispersal
between ecoregions might be limited because large portions of the
surrounding matrix are unsuitable for forest species. Differences in
vagility might cause the observed guild patterns. The diversity be-
tween ecoregions increased from fresh-wood species to rotten-
wood species to fungi-associated species; the opposite was true
for a diversity. This indicates that a fresh-wood colonizer has to
be slightly more mobile (Müller and Goßner, 2007; Speight et al.,
2003) because it has less time to colonize the host substrate than
a colonizer of a decaying tree or of fungi, which are more concen-
trated over a longer period in their decaying substratum and are
more sensitive to forest fragmentation (Komonen et al., 2000).
However, even for a fungi colonizer, distances of kilometres can
be bridged (Jonsson and Nordlander, 2005; Ranius, 2006), but the
dispersal of saproxylic beetles over larger distances, e.g. from eco-
region to ecoregion, has not been studied. Our results provide some
indication that dispersal limitation might not be the most impor-
tant reason for b diversity because the scale between our forest
sites, which were also separated by non-forest land, was of minor
importance.
A second explanation for the importance of b diversity between
ecoregions may be differences in tree species. Host species differ
greatly in insect species richness, and therefore, the observed
diversity patterns might be biased because of the different tree
species composition in the different ecoregions. For example, oak
has the most insect-species-rich fauna among the tree species of
Central Europe (Brändle and Brandl, 2001; Stork et al., 2001). How-
ever, the most-abundant tree species in all ecoregions were sam-
pled, and oak was sampled in most ecoregions. Furthermore,
only a few saproxylic beetles in Central Europe are specialised on
a single tree species or genus (for an example of a saproxylic family
rich in species, see Cerambycidae in Bense, 1995). Therefore, a bias
caused by effects of different tree species sampled in the ecore-
gions is expected to be low, although isolated host trees might
have an effect (Müller and Goßner, 2007).
The third explanation for the importance of b diversity between
ecoregions might be different soil conditions, climates, and degrees
of naturalness owing to biogeographic and land-use histories
(Rüther and Walentowski, 2008). In our data, we found some indi-
cations for these processes by the lower number of species in the
Tertiäres Hügelland ecoregion. In this landscape, the soil is highly
productive, and much of the naturally occurring broadleaf forests
have therefore been replaced by large-scale spruce plantations.
To test the influence of forest sites with altered tree species
composition on b diversity pattern, we analysed also only those
sites with a close-to-natural tree species composition. The critical
631
diversity patterns obtained were similar to those obtained when
all sites were analysed (see Supplementary material). Our results
indicate that the differences in overall c diversity were neither
caused simply by fragmentation nor by tree species composition.
More likely the different soil and climatic conditions, differences
in tree species composition, and site history together form new
habitats for new species in the ecoregions. These findings might
be particularly important for the development of state-wide strat-
egies. Most recent saproxylic beetle studies, which were restricted
to local sites, have pointed out the importance of fragmentation
(Brunet and Isacsson, 2009; Goßner et al., 2008; Müller and
Goßner, 2007) or tree species composition (Jonsell et al., 2007)
on the local level. As soon as studies on different spatial scales, like
the one presented here, reveal the high importance of a more com-
prehensive, general habitat type, factors can be identified for the
sustainability of the biodiversity of saproxylic beetles on the land-
scape scale. Since most forested landscapes in temperate zones are
highly fragmented by silviculture, it is difficult to quantify the pro-
portion of b-5 diversity under natural conditions in an almost com-
pletely covered forested landscape. It might be assumed that a
better exchange of species and more homogenous conditions in
natural forests on a broader scale would lead to a lower b-5 diver-
sity. Differences in climate conditions and in the dominance of for-
est vegetation types, however, would clearly be obvious even
under natural conditions. However, even if it is difficult to assess
this effect, conservation planning nowadays has to deal with the
existing highly fragmented landscapes.
This is also supported by our findings on the small scale, with
important b diversity among stands within forest sites. In contrast,
a generally lower proportion of b diversity was found among
microhabitats (among traps), vertical strata (understory/canopy),
and among forest sites within an ecoregion. The importance of
the diversity between stands can be reasoned by different condi-
tions in the stand structure, tree species composition, amount of
dead wood, and the history of the stands, all of which influence
saproxylic communities (Goßner et al., 2008; Müller and Bussler,
2008; Rüther and Walentowski, 2008).
The low a diversity within traps and the low b diversity among
traps and vertical strata indicate that dispersal between microsites
is not limited for most species in our study. We have to keep in
mind that we trapped flying saproxylic beetles, for which dispersal
is not limited within distances <100 m, not even for specialists
(Müller and Goßner, 2007; Ranius, 2006; Schiegg, 2000). Conse-
quently, for most species, movement from the ground to the can-
opy, by flying or by climbing the stems (Goßner, 2004), should
not be limited, which results in a low proportion of b diversity
among strata. These results are confirmed by the additive parti-
tioning of diversity of Lepidoptera and Coleoptera in temperate for-
ests in Japan (Hirao et al., 2007), even if several other studies have
shown stratification of arthropod assemblages in temperate forests
(Goßner et al., 2009; Gruppe et al., 2008; Stork et al., 2001). The
proportion of a diversity within traps being consistently signifi-
cantly lower than expected by chance might also be explained by
the general rule of inflation of b diversity and a decrease of a diver-
sity caused by the aggregation of individuals of one species (Veech,
2005). Particularly saproxylic beetles aggregate, especially on the
local scale because of their specific host substratum and mating
sites.
4.2. Are species in low numbers threatened species?
Gering et al. (2003) used the density of beetle species in fogging
samples to classify ‘‘rare” and ‘‘common” species, but they were
uncertain whether this definition is a sampling artefact or
biologically meaningful. However, such a posteriori classification
from data always comprises somewhat circular reasoning. The
632 J. Müller, M.M. Goßner / Biological Conservation 143 (2010) 625–633
current red list of Bavaria, which is based on broad expertise
throughout the country and on many studies in different types of
forests (Schmidl et al., 2003), enabled us to use an a priori classifi-
cation of non-threatened species as ‘‘common” and threatened
species as ‘‘rare”. When we compared the results of both ap-
proaches in our study, the diversity partitioning pattern of species
defined a priori as threatened was similar to the pattern of species
with low density in the samples. This supports the widely held be-
lief that most rare species in the samples are threatened within a
region. For the more abundant and common species, we found a
distinct pattern with some overestimation of a diversity for ‘‘com-
mon” species using the abundant species from sampling data. This
indicates that in each type of standardized sampling, a fixed pro-
portion of species, even if common in a state, have low densities
down to singletons. However, our results support the general find-
ings of Gering et al. (2003) that the richness of rare species is influ-
enced most by b diversity and that threatened species mirror the
diversity pattern of the entire community (Fig. 5).
4.3. Monitoring
Even with the comprehensive literature on beetle species in
Central Europe, some families of beetles, e.g. Cryptophagidae and
Staphylinidae, can only be determined by a few experts. Therefore,
large-scale studies are restricted by financial constraints to a sub-
set of species (Duelli and Obrist, 1998; Ferris and Humphrey,
1999). Our analysis showed that the subset of ‘‘monitoring species”
for Germany as proposed by Schmidl and Bußler (2004) mirrored
the diversity pattern of the entire community, and can therefore
be used in monitoring studies.
Our data from near ground and the canopy additionally allowed
us to test the importance of the third dimension (for European for-
ests, see Floren and Schmidl, 2008) for understanding the impor-
tant diversity scales. Our results clearly indicated that the b
diversity estimates of near ground and canopy are similar. There-
fore, the canopy could be neglected in monitoring projects for
beta-diversity assessments between sites, without a substantial
loss of information. However, for measuring the effects of local
management within sites, it might be very important to consider
the canopy, because the canopy composition differs from lower
stratum, as indicated by Goßner (2009) and Su and Woods
(2001). In both of these studies, clear differences in the density
and diversity of the canopy and near-ground strata were found
in different structured forest stands. These studies, however, fo-
cused only on single sites in contrast to our broad-scale approach.
In general, our study demonstrates that the broadly applied
monitoring of biodiversity of saproxylic organisms using indirect
assessments of indicators/proxies such as dead wood amount
(Bütler and Schlaepfer, 2004) or tree age (Moning and Müller,
2009), show that such approaches obscure important scales and
factors for diversity because they neglect the complex interaction
of habitat types, climate and species distribution. Therefore, we
strongly recommend a species-based concept in conservation plan-
ning and monitoring.
5. Conclusion
Our findings are relevant for designing networks of areas with
conservation priority on local and regional scales. Conservationists
still debate whether it is preferable to design reserves as a single
large cluster or as several small (SLOSS) patches (reviewed by
Shafer, 2001). Our additive partitioning of saproxylic beetle diver-
sity generally supports the establishment of conservation areas
that represent as many ecoregions as possible on a regional,
state-wide scale, similar to that attempted within the network of
Natura 2000. We in particular suggest that this network be en-
larged, especially in underrepresented ecoregions. On the local
scale, the second major part of b diversity was between stands,
which favours the protection of new reserves and forest stands
with conservation-oriented forestry rather than to enlarge existing
reserves.
The widely applied conservation practice of focusing on threa-
tened species is also supported by our results. The important spa-
tial scale for threatened species (ecoregions and stands) also
encompasses the spatial scales that appear to be important for
the richness and diversity of common species. This result is note-
worthy because conservation management strategies could merge
the conservation of rare species richness with the maintenance of
forest functions driven more by common insect species. For state-
wide monitoring of the prominent group of saproxylic beetles, a
limitation to ‘‘monitoring species” and to near-ground strata seems
sufficient to represent the general patterns of biodiversity. Finally,
we want to point out that information about the spatial scales of
biodiversity should not be contemplated as an alternative for re-
quired species-habitat relationships, as e.g. rotholes (Bussler and
Müller, 2009), but rather as a valuable technique that can help to
develop landscape-wide effective strategies. To increase our
understanding, we clearly recommend diversity partitioning of
other diverse species groups in forest conservation.
Acknowledgements
We are grateful to Bernhard Förster for help with the database,
to the Bavarian State Institute of Forestry for providing data and to
Karen A. Brune for linguistic revision of the manuscript.
Appendix A. Supplementary material
Supplementary data associated with this article can be found, in
the online version, at doi:10.1016/j.biocon.2009.11.027.
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