The influence of species traits and q-metrics on scale-specific b-diversity components of arthropod communities of temperate forests
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Landscape Ecol (2011) 26:411–424
DOI 10.1007/s10980-010-9568-9
RESEARCH ARTICLE
The influence of species traits and q-metrics on scale-specific
b-diversity components of arthropod communities
of temperate forests
Martin M. Gossner • Jörg Müller
Received: 14 April 2010 / Accepted: 23 December 2010 / Published online: 8 January 2011
Ó Springer Science+Business Media B.V. 2011
Abstract Protection of biodiversity and ecosystem ecoregion level than generalist species. Differences in
functions requires a better understanding of spatial the proportion of b-diversity between functional
diversity. Here we studied diversity patterns of true guilds increased with increasing weighting of abun-
bugs and saproxylic beetles, sampled in 28 forest dant species. The b-diversity patterns based on body
stands of southern Germany, using a hierarchical size and host specificity were similar for true bugs,
nested design of five increasingly broader spatial but partly contrasting for saproxylic beetles. Our
levels: trap location, stratum, forest stand, forest site, results suggest that (1) future conservation schemes
and ecoregion. We predicted that: (1) for large body- should focus on establishing new conservation sites
sized species (as a surrogate for highly mobile in new ecoregions, rather than on enlarging existing
species) and host generalist species (low host spec- protected areas; (2) host specificity might be a more
ificity), the proportion of b-diversity decreases from meaningful trait than body size to be considered in
small to large spatial scales; and (2) the differences biodiversity studies; and (3) common conservation
between trait-based functional guilds in the propor- approaches restricted to only large, conspicuous, but
tion of b-diversity increase with increasing weighting rare species might result in a mismatch of important
of more-abundant species. Our results indicated that biodiversity scales.
the ecoregion level is the most important diversity
scale for both taxa and among functional guilds Keywords Spatial scale Multiplicative diversity
sampled, followed by the forest stand level. Special- partitioning Body size Host specificity Host niche
ized species were more strongly affected on the breadth Ecosystem function
Electronic supplementary material The online version of
Introduction
this article (doi:10.1007/s10980-010-9568-9) contains
supplementary material, which is available to authorized users. Reducing the loss of global biodiversity (Balmford
et al. 2005a, b) and maintaining ecosystem functions
M. M. Gossner (&)
Institute of Ecology, Friedrich Schiller University, require a better understanding of a- and b-diversity in
Dornburger Str. 159, 07743 Jena, Germany relation to species traits across spatial scales, from
e-mail: martin.gossner@uni-jena.de both ecological and economical points of view
(Zavaleta and Hulvey 2004; Bunker et al. 2005;
J. Müller
Bavarian Forest National Park, Freyunger Str. 2, 94481 Hooper et al. 2005; Spehn et al. 2005; Cardinale et al.
Grafenau, Germany 2006; McIntyre et al. 2007). Macroecological
123412 Landscape Ecol (2011) 26:411–424
biodiversity patterns on the global scale have been guilds differ with spatial scales, as demonstrated for
recognized, but our ecological understanding of the forest moths of different body size and niche
dynamics of such patterns is still limited (Brown et al. specificity.
2004; Qian et al. 2005; Qian and Ricklefs 2007, Species traits and spatial scale are also related with
2008). At regional scales, recent studies have sug- regard to habitat fragmentation, with highly disper-
gested that species turnover (b-diversity) is more sive species being able to shift more easily to and
important for developing conservation strategies than from distant forest patches across a region than less-
comparing a-diversity (Basset et al. 2008; Müller and dispersive species (Nekola and White 1999; Didham
Brandl 2009). In general, species composition is and Fagan 2004). Investigations on the two traits
strongly correlated with geographical distance mobility and host specificity have revealed that host
(Harrison et al. 1992; Rosenzweig 1995; for arthro- specialists are less mobile and more closely related to
pods, e.g., Tylianakis et al. 2006), but it is far from specific habitat conditions. Therefore, the species
clear how increasing the spatial scale really translates turnover might in general be weaker for species with
into increasing species turnovers. This is enforced by low host specificity than for species with high host
limitation of most studies to only a few spatial or specificity (Komonen et al. 2004; Hirao et al. 2007).
temporal scales (e.g., Hirao et al. 2007). Partitioning c-diversity according to spatial scales
Several spatial scales for forest-dwelling arthro- (Whittaker 1960), from the stratum of a single tree or
pods can be considered: the local scales of a stand, local stands up to landscapes, has been developed
i.e., the trap location as the sampling unit in a tree and further (Lande 1996; Veech et al. 2002). Researchers
stratum; and larger scales, i.e., stands, forests, and have applied this method to several taxonomic groups
ecoregions (Gering et al. 2003). Most previous of insects, mainly moths and beetles (Summerville
studies of forest arthropods focused only on one or and Crist 2002, 2003; Gering et al. 2003; Summer-
a few of these spatial scales because of restricted ville et al. 2003; Veech 2005; Müller and Gossner
sampling methods (e.g., on the stand, forest, and 2010; Röder et al. 2010), but only a few studies have
ecoregion levels in the studies of Summerville et al. considered species traits (Summerville and Crist
2003, 2006), although a few also considered the tree 2002; Summerville et al. 2006; Röder et al. 2010).
level (Gering and Crist 2002; Gering et al. 2003). The recent introduction of a general q-metric
Furthermore, the conspicuous vertical gradients of based on multiplicative partitioning, however, has
forests in structure, biomass, light, and temperature improved the possibilities for calculating b-compo-
were widely neglected until fairly recently (Parker nents along a continuous gradient of increasing
1997; Basset et al. 2003; Horchler and Morawetz weight of abundant species. This could be important
2008; Tal et al. 2008). Thus, although there might be for biodiversity conservation because conserving
only minor shifts in species composition across abundant species could be critical for conserving
forests of an entire ecoregion, there could be major ecosystem functions (Taylor et al. 2006; Gaston
shifts from the forest canopy to the forest floor within 2010).
forests, as shown for butterflies and beetles in a cool- Here we collected true bugs and saproxylic beetles
temperate deciduous forest (Hirao et al. 2007). using flight-interception traps to test the following
Studies have increasingly linked biodiversity with a priori predictions for trait- and scale-related
functional diversity (Ernst and Rödel 2005; Balva- dependency of diversity in forest insect communities:
nera et al. 2006; Klein et al. 2008), but how species (1) for large body-sized species (highly mobile) and
traits and different scales interact is still poorly generalist species (low host specificity), the propor-
understood. Recent progress in describing macroeco- tion of b-diversity will decrease from small (trap
logical patterns have revealed body size (as an location, vertical stratum) to intermediate (stand,
indirect measure of dispersal ability) and host spec- forest) to large (ecoregion) spatial scales, and the
ificity as major determinants of species bionomics opposite pattern will be found for species of low
and geographical distribution (Blackburn and Gaston mobility (small body size) and high host specificity,
2003; Komonen et al. 2004; Bertheau et al. 2010; with intermediate patterns for species of medium
Chown and Gaston 2010). Furthermore, Summerville body size (intermediate mobility) and host specific-
et al. (2006) described how trait-based functional ity; and (2) the differences between functional guilds
123Landscape Ecol (2011) 26:411–424 413
in the proportion of b-diversity on different spatial restrictions) representing the dominant tree species in
scales will increase with increasing weighting of the overstorey were selected randomly. Traps were
more-abundant species. installed pair-wise in the center of the selected tree
crowns (16–33 m) and near the ground (1.5 m). All
adult true bugs and saproxylic beetles were identified
Methods to the species level (for details see supplement S2).
Among beetles, saproxylic beetles were defined
Study sites and sampling design according to the definitions of Speight (1989).
True bugs and saproxylic beetles were sampled in Trait characterization of species
mature forest stands (age [80 years) in southern
Germany in different projects from 1996 to 2007 Species traits were characterized according to the
following a standardized protocol. We tested and data given in Wachmann et al. (2004–2008) and
standardized the collecting effort for each trap during Böhme (2005). Fortunately, the ecology of saproxylic
the first 2 years of the study and then used the beetles and true bugs is well studied in Germany,
standardization in all following projects (for more resulting in comprehensive data that allow reliable
details and validation on sampling design and deter- trait-based statistical analysis. By using a rougher
mination, see supplementary material S1 and Müller classification, uncertainties in the trait characteriza-
and Gossner 2010). tion of a few species could be circumvented. Previous
We created a hierarchically nested data matrix that studies have shown that body size is correlated with
included categories for the main ecological forest species dispersal ability (for insects, e.g., Brändle
types of the area. The five hierarchical levels et al. 2002). Therefore, we used the body size of the
corresponded to the following spatial scales: ecore- species given in literature (beetles: Freude et al.
gion, forest (connected forest area [ 500 ha), stand 1964–1983, true bugs: Wachmann et al. 2004–2008)
(5–70 ha), two strata within the stand (near ground as a surrogate for dispersal ability. A frequency
and canopy), and the trap locations within the stratum distribution of the body size of all saproxylic beetle
of a stand (for details see supplementary material S1). species and of all true bug species of Germany was
Henceforth, for simplification we use the term ‘traps’ plotted a priori and divided into three categories of
as the smallest spatial scale. The highest level and equal number of species (Fig. 1). This resulted in
therefore the broadest spatial scale was represented body size classes of saproxylic beetles of small:
by five ecoregions (Mainfränkische Platte, Fränki- \2.5 mm, medium: 2.5–6 mm, and large: [6 mm
scher Keuper, Frankenalb, Bayerischer Wald, and and of true bugs of small: \4 mm, medium:
Tertiäres Hügelland) containing mixed oak forests, 4–6.2 mm, and large: [6.2 mm.
beech/oak-dominated forests, spruce-dominated plan- Host specificity of each arthropod taxon was
tations, mixed montane forests, and high montane classified into three categories (for a complete list,
spruce forests (Table S1; Müller and Gossner 2010). see Supplement S5). Monophagous species of phy-
The classification of ecoregion (see also Fig. S1) is tophagous and zoophytophagous true bugs that feed
based on similar geological and climatic conditions on plant species of one genus only were classified as
for forest growth (Walentowski et al. 2006). species with high host specificity. Oligophagous
We used flight-interception traps consisting of a species of true bugs that feed on more than one plant
crossed pair of transparent plastic shields (40 9 genus up to five plant families were classified as
60 cm) with funnels opening into sampling jars at the species with medium host specificity. Polyphagous
bottom and at the top. Insects were trapped passively species of true bugs that feed on more than five plant
over one entire vegetation period (April to October) families were classified as species with low host
(see Gossner 2008). Hence, each sample represents specificity. Predacious true bugs species that show a
the community of the whole vegetation period and narrow secondary host specificity by feeding only on
owing to very low insect activities during the winter, plant species of one genus (e.g., Deraeocoris annul-
also almost the entire community. In each stand, five ipes feeds on Aphididae, exclusively on Larix
trees (in two forests, only three trees owing to project decidua), or a broader secondary host specificity by
123414 Landscape Ecol (2011) 26:411–424
Fig. 1 Body-size distribution of all true bug (a) and saproxylic beetle (b) species recorded in Germany. Each species was classified
into one of three body size classes of approximately equal number of species. Class borders are indicated by dashed lines
feeding only on one to five plant families (e.g., using true diversities and q-metric (see below), which
Anthocoris confusus feeds on Homoptera, mainly on are implemented in the approach advocated by Jost
Fagaceae, more rarely on Aceraceae, Tiliaceae, (2006, 2007), we decided to use the multiplicative
Oleaceae, Salicaceae) were classified as species with approach, based on Whittaker’s (1960) formula:
a high and a medium host specificity, respectively.
c ¼ a1 ðwithin trapÞ b1 ðamong trapsÞ
Predacious species of true bugs that feed on more
than five plant species were classified as species with b2 ðamong strataÞ b3 ðamong standsÞ
low host specificity. Saproxylic beetles were classi- b4 ðamong forestsÞ b5 ðamong ecoregionsÞ:
fied into the following categories of host specificity,
irrespective of trophic level: high: feeding on species When using ‘‘numbers equivalent’’ (effective number
of one tree genus; medium: feeding on either broad- of species), which has the properties expected from a
leaved or coniferous trees; low: feeding on broad- true diversity measure (Jost 2007), index-independent
leaved and coniferous trees. formulas can be derived in which an exponent value
q indicates the order of the diversity measure
(Keylock 2005). By using this q-metric (Jost 2007),
Data analysis
functional trait patterns can be analyzed with contin-
uous weighting from rare (low q value) to abundant
c-diversity can be partitioned into a- and b-diversity
(high q value) species (Keylock 2005; Jost 2007).
components either additively (Veech et al. 2002), as
For all species i = 1–S, a-diversity, c-diversity, and
done in most previous studies, or multiplicatively
b-diversity are calculated (see Jost 2007) according to:
(Whittaker 1960; Jost 2007). Jost and colleagues (Jost
" #1=ð1qÞ
2006, 2007; Jost et al. 2010) recommend the use the XN X
q q
multiplicative approach because of the dependency of Da ¼ pi wj ð1Þ
b-diversity on the a-diversity in additive partitioning. j¼1
However, as demonstrated in a Forum in Ecology " #1=ð1qÞ
S X
X
(Veech and Crist 2010), neither the additive nor q
Dc ¼ pij wqj ð2Þ
the multiplicative approach is able to produce a i¼1
b-diversity statistically independent of the a-diver- q
Db ¼q Dc =q Da ð3Þ
sity, as suggested by Jost (2006, 2007), Tuomisto and
Ruokolainen (2006), and Jost et al. (2010). After where pi is the proportional abundance of species i in
inspecting all arguments offered in this forum and sample j, wj is the weight of the sample (in our case
previous publications and owing to the advantages of weights are equal = 1/N), and q is the q value.
123Landscape Ecol (2011) 26:411–424 415
To analyze how differences in the abundance of opposite pattern was observed for saproxylic beetle
species affect the proportion of b-diversity of species species (high host specificity: 89; low host specificity:
of different species traits, we used q values from 0 to 61).
4 in steps of 0.5. All q values \ 1 are disproportion- In general, the smallest (among traps) and largest
ately sensitive to rare species, and q values [ 1 are (among ecoregions) spatial scale contributed most to
disproportionately sensitive to more-abundant spe- the total b-diversity when either body size or host
cies. q = 0 corresponds to species richness; specificity was considered, followed by the among-
q = 0.999 (and not q = 1, which would require stand scale (Figs. 2, 3, 4). The among-stand scale was
division by zero) produces the widely distributed related more to species turnover for true bugs than for
Shannon diversity; and q = 2 corresponds to the saproxylic beetles. Species turnover at the among-
often used Simpson diversity measures. We multipli- forests and among-strata scales was comparatively
catively partitioned the community divided into low.
trait-based functional guilds using the software
PARTITION 3.0 (Veech and Crist 2009) without
sample weighting. Because we focused on the
differences in the proportions of the b-diversity
(a) True bugs Saproxylic beetles
levels, only these results are given. For comparative 100 100
reasons, additive partitioning results are also pre- 80
80
sented in Supplement S4, including the a-diversity of
Diversity
Diversity
the smallest spatial scale. In contrast to multiplicative 60 60
partitions, which show effective numbers of species 40 40
(for all values of q), additive partitions show the
absolute number of species (at least in the case where 20 20
q = 0). Because multiplicative and additive parti- 0 0
tions involve calculations of interdependent compo- small medium large small medium large
Body size Body size
nents of a- and b-diversity across scales and because
incomplete sampling at each scale is assumed, null (b) True bugs Saproxylic beetles
100 100
randomization tests were applied (Veech and Crist
2009). Details on the null randomization tests are 80 80
given in Supplement S4.
Diversity
Diversity
60 60
40 40
Results
20 20
General patterns 0 0
high medium low high medium low
Host specificity Host specificity
Our final data set consisted of 147 true bug species
(5,083 individuals) and 470 saproxylic beetle species β5 (among ecoregions) β2 (among strata)
β4 (among forests) β1 (among 'traps')
(23,985 individuals). The distribution of body sizes of β3 (among stands)
true bugs was slightly shifted toward larger species,
and that of saproxylic beetles was comparable to that Fig. 2 Multiplicative diversity partitioning (without sample
found in Germany (see Fig. S3), i.e., we observed weighting) of species of different body size (a) and host
specificity (b). The percentage of diversity (q-metrics: q = 2)
more large (68) than medium (41) or small (38)
explained by b-components on five spatial scales is shown:
species of true bugs, and most saproxylic beetle among trap locations (‘traps’) of one stratum within a stand and
species were of medium size (168), followed by small strata (low spatial scale, white bars), forest stands and forests
(163) and large (139) sizes. Most true bug (70) and sites (medium spatial scale, gray bars), and ecoregions (large
spatial scale, black bars). Connecting lines among bars
saproxylic beetle species (320) had medium host
separate the three different spatial scales. The contributions
specificity. More true bug species had low host to the total c-diversity for each scale were determined using
specificity (51) than high host specificity (26); the Whittaker’s multiplicative formula
123416 Landscape Ecol (2011) 26:411–424
Fig. 3 Multiplicative
Body size
diversity partitioning small medium large
(without sample weighting)
of species of different body True bugs Saproxylic beetles
size. The percentage of β-diversity among ecoregions β-diversity among ecoregions
Proportion of β-diversity
Proportion of β-diversity
diversity explained by 0.38 0.55
0.36 0.50
b-components on five
0.34 0.45
spatial scales is shown, with 0.32
a decreasing spatial scale 0.40
0.30
from top to bottom: 0.35
0.28
ecoregions (large spatial 0.26 0.30
scale), forests sites and 0.24 0.25
forest stands (medium 0.22 0.20
0 1 2 3 4 0 1 2 3 4
spatial scale), strata and trap
location (‘traps’) (low q-value q-value
spatial scale). The
β-diversity among forests β-diversity among forests
Proportion of β-diversity
Proportion of β-diversity
contributions to the total 0.16 0.16
c-diversity for each scale
0.15 0.15
were determined using
q-metrics (see Jost 2007) 0.14 0.14
and Whittaker’s 0.13 0.13
multiplicative formula.
With increasing q value, the 0.12 0.12
increasing influence on 0.11 0.11
abundant species is given in 0 1 2 3 4 0 1 2 3 4
the calculation of diversity, q-value q-value
calculated from the relative
abundance of species in the β-diversity among stands β-diversity among stands
Proportion of β-diversity
Proportion of β-diversity
0.24 0.22
samples. q = 0 corresponds
to the calculation of species 0.22 0.20
richness, q = 1 (0.999 was 0.20
0.18
used) corresponds to the 0.16
calculation of Shannon 0.18
0.14
diversity. A total of 147 true 0.16 0.12
bug species and 470
0.14 0.10
saproxylic beetle species 0 1 2 3 4 0 1 2 3 4
were sampled (small body
q-value q-value
size: 38/163; medium body
size: 41/168; large body β-diversity among strata β-diversity among strata
Proportion of β-diversity
Proportion of β-diversity
size: 68/139). Body size 0.19 0.19
0.18
classes were defined a priori 0.18
0.17
based on size distributions 0.16
0.17
of all species recorded in 0.15
Germany (see Fig. S3) 0.16 0.14
0.13
0.15
0.12
0.14 0.11
0 1 2 3 4 0 1 2 3 4
q-value q-value
β-diversity among 'traps' β-diversity among 'traps'
Proportion of β-diversity
Proportion of β-diversity
0.25 0.30
0.24 0.28
0.23 0.26
0.22 0.24
0.22
0.21
0.20
0.20 0.18
0.19 0.16
0.18 0.14
0.17 0.12
0 1 2 3 4 0 1 2 3 4
q-value q-value
123Landscape Ecol (2011) 26:411–424 417
Fig. 4 Multiplicative
Host specificity
diversity partitioning
high medium low
(without sample weighting)
of species of different host True bugs Saproxylic beetles
specificity. For β-diversity among ecoregions β-diversity among ecoregions
Proportion of β-diversity
Proportion of β-diversity
methodological details, see 0.45 0.50
Fig. 3. A total of 147 true 0.40 0.45
bug species and 470 0.35 0.40
saproxylic beetle species
0.30 0.35
were sampled (high host
0.25 0.30
specificity: 26/89; medium
host specificity: 70/320; low 0.20 0.25
host specificity: 51/61). For 0.15 0.20
0 1 2 3 4 0 1 2 3 4
the classification of host
specificity, see ‘‘Methods’’ q-value q-value
β-diversity among forests β-diversity among forests
Proportion of β-diversity
Proportion of β-diversity
0.15 0.16
0.15
0.14
0.14
0.13
0.13
0.12
0.12 0.11
0 1 2 3 4 0 1 2 3 4
q-value q-value
β-diversity among stands β-diversity among stands
Proportion of β-diversity
Proportion of β-diversity
0.24 0.22
0.22 0.20
0.18
0.20
0.16
0.18
0.14
0.16 0.12
0.14 0.10
0 1 2 3 4 0 1 2 3 4
q-value q-value
β-diversity among strata β-diversity among strata
Proportion of β-diversity
Proportion of β-diversity
0.18 0.19
0.18
0.17
0.17
0.16 0.16
0.15 0.15
0.14
0.14
0.13
0.13 0.12
0 1 2 3 4 0 1 2 3 4
q-value q-value
β-diversity among 'traps' β-diversity among 'traps'
Proportion of β-diversity
Proportion of β-diversity
0.28 0.28
0.26 0.26
0.24 0.24
0.22 0.22
0.20 0.20
0.18 0.18
0.16 0.16
0.14 0.14
0 1 2 3 4 0 1 2 3 4
q-value q-value
123418 Landscape Ecol (2011) 26:411–424
Body size and b-diversity proportions of dominant species, whereas smaller spatial scales
were more important for the turnover of rare species.
The body-size-related b-diversity patterns differed However, we need to consider that by weighting rare
between taxa and trait-based functional guilds species, the proportion of b-diversity on the higher
(Fig. 2). For large-sized species of both taxa, the spatial scale did not differ much from that on the
smallest spatial scales (among traps and strata) were smallest scale. In contrast, when abundant species
most important and the largest (among ecoregions) were weighted, these two scales clearly differed.
were least important. Small- and medium-sized The overall differences between trait-based func-
species of true bugs revealed a similar pattern with tional guilds of saproxylic beetles were relatively
almost equal importance of the three coarse spatial small when rare species were weighted and increased
scales, whereas the highest spatial scale seemed to be with increasing weighting of more-abundant species;
much more important for medium-sized saproxylic this was observed for true bugs almost only at
beetle species and least important for small-sized intermediate spatial scales. In several cases, the
saproxylic beetle species. Species turnover on relative influence of scale on species turnover asso-
medium spatial scales (among stands, among forests) ciated with different body-size classes changed with
was comparatively low for both taxa. relative abundance (Fig. 3). For example, the pro-
portion of among-ecoregion b-diversity of saproxylic
Host specificity and b-diversity proportions beetles was highest for small-sized species when
more weight was given to rare species, and highest
Host-specificity patterns of the saproxylic beetles and for medium-sized species when more weight was
true bugs were more similar than their body-size given to abundant species. In contrast, the among-trap
patterns, e.g., the relative importance of the ecoregion b-diversity was higher for medium-sized species
scale decreased more or less linearly from high to low when more weight was given to rare species and
host specificity for both taxa, but from small to large higher for small-sized species when more weight was
body size only in true bugs; saproxylic beetles showed given to the most-abundant species. In the host
a clear humped-shaped pattern (Fig. 2). The among- specificity patterns, the b-diversity of the different
ecoregions scale was related most to species turnover functional guilds also changed in a few cases (i.e.,
for specialized species (high host specificity), whereas true bugs at the among-forests level) from being more
all other scales were related more to species turnover important for rare species to being more important for
in species with medium and low host specificity. We common species, but the patterns differed less than
observed only slight differences between saproxylic body-size patterns (Fig. 4).
beetle species with medium and low host specificity The body-size and host specificity patterns of true
(Fig. 2). Mainly the among-stands and among-traps bugs were similar, whereas those of saproxylic beetles
scales contributed most to species turnover for true differed, sometimes even completely, especially at the
bugs with low host specificity; the among-ecoregions, ecoregion, forest, and stratum spatial scales. When we
among-forests, and among-strata scales contributed focused on the relatively abundant species, the
most to species turnover for species with medium host ecoregion scale was much more important for large-
specificity (Fig. 2). sized than for medium- or small-sized saproxylic
beetle species, whereas this scale was most important
The influence of weighting rare and common for host specialists. The proportion of species turnover
species observed at the among-forests and among-strata
scales, in contrast, was highest for large-sized species
With increasing weighting of more-abundant species, and for species of intermediate or large host specific-
the among-ecoregions scale (and partly the among- ity. Different patterns of body size and host specificity
forests scale) accounted for more species turnover, of true bugs was observed only at the among-traps
and the among-stands and among-traps scales (and level. This level was more important for small-sized
partly the among-strata scale) contributed less species species than for medium-sized species, but was more
turnover (Figs. 3, 4). This means that larger spatial important for species with medium host specificity
scales were relatively more important for the turnover than with high host specificity.
123Landscape Ecol (2011) 26:411–424 419
Discussion stands than for saproxylic beetles, which indicated
that suitable habitats for saproxylic beetles might be
General patterns more evenly distributed among stands than those for
true bugs; true bugs living mainly on green trees may
We found that the smallest spatial scale (among- be more specialized on specific habitat conditions of
traps, but not among-strata) and largest spatial scale their host trees, which makes the insects in general
(among-ecoregions) contributed most to the total less mobile (Brändle et al. 2002; Komonen et al.
b-diversity when either body size or host specificity 2004), and vice versa, saproxylic beetles occurring in
was considered. This indicates that species turnover ephemeral habitats have to be more successful at
occurred mainly on a horizontal organization level of finding suitable conditions for their larvae on larger
trees within forest stands and among ecoregions. scales (Müller and Gossner 2007).
Considering insect species, it seems that suitable
habitats, such as dead wood structures or host plants, Functional traits and b-diversity proportions
are distributed in horizontal patches mainly within
forest stands, which leads to the aggregation of Our hypothesis that the proportion of b-diversity of
saproxylic beetles and true bugs (Sobek et al. 2009; highly mobile and generalist species decreases from a
Ylisirnio et al. 2009). Although some vertical small to a large spatial scale was confirmed by the
stratification of the fauna in Central European forests results obtained for true bugs, with a relatively higher
has been documented (Gruppe et al. 2008; Gossner turnover of small and specialized species at the
2009), when more than two spatial levels (canopy and ecoregion scale. Species turnover of more generalist
near ground) of diversity were considered, vertical species occurred mainly on smaller spatial scales,
species turnover within forest stands appeared to be which suggested a patchy distribution based upon the
lower than horizontal turnover. One of the main patchy distribution of their host plants (Ribeiro et al.
explanations for this seems the rareness of ‘real’ 2003; Summerville et al. 2003).
canopy species in Central European temperate for- In contrast, results obtained with saproxylic bee-
ests, mainly owing to decreased habitat diversity in tles supported our hypothesis only when host spec-
the upper stratum of temperate trees (for discussion, ificity was considered. The relatively high species
see Müller et al. 2008). turnover of specialized species on the ecoregion level
We previously described high species turnover and low species turnover on small spatial scales
among ecoregions (Müller and Gossner 2010) and might reflect a patchy distribution of suitable
related it to three main factors: (1) limited dispersal resources on a larger scale. Moreover, several
between ecoregions because of highly unsuitable specialized saproxylic species might exhibit a strong
landscape connectivity, (2) differences in tree-species relationship to habitat continuity (Müller et al. 2005)
composition leading to highly varying insect species and habitat conditions, combined with a low dispersal
richness owing to tree species specificities, and (3) willingness (Jonsson 2000; Ranius and Heding 2001),
different soil conditions, climates, and degrees of which may lead to higher turnover between commu-
naturalness reflecting biogeographic and land-use nities in different ecoregions. Unfortunately, we do
histories. Species turnover among ecoregions is not yet have physical measurements of dispersal
perhaps driven by these environmental and habitat ability and willingness of whole insect communities
conditions and factors linked to dispersal (barriers, to disperse, and therefore surrogates have to be used
movement ability, and behavior) (Soininen et al. (Komonen et al. 2004).
2007), as has been demonstrated for leaf beetles
(Baselga and Jimenez-Valverde 2007), butterflies Body size versus host specificity
(Dover and Settele 2009), and geometrid moths
(Beck and Khen 2007). We found that the patterns of both taxa were clearly
Medium spatial scales (among-forests, among- more apparent and consistent with our hypothesis
stands) contributed substantially to total diversity, but when we used host specificity as an indirect measure
to a lesser extent than small and large spatial scales. of specialization than when we used body size as a
Species turnover was higher for true bugs among measure of dispersal ability. The relationship
123420 Landscape Ecol (2011) 26:411–424
between body size and host range, however, seems to might have additionally affected the diversity parti-
be very robust for European Heteroptera (Brändle tioning pattern, as has been shown for Lepidoptera
et al. 2000), and 75% of 24 such studies showed a (Lindstrom et al. 1994) in a temperate deciduous
positive correlation (Loder et al. 1998). Most of these forest of North America (Summerville et al. 2006).
studies were on butterflies and moths, but a positive To obtain better results, we clearly need more
correlation was observed also in other phytophagous progress in developing better functional traits that
insect communities, e.g., of leaf beetles and weevils mirror the behavior of species, even if it seemed
(Loder et al. 1998) and of tropical sap-suckers and advantageous to use a measured and well-known
leaf-chewers (Novotny and Basset 1999). A positive parameter such as body size instead of expert
correlation between body size and host range has assessments for dispersal ability (Komonen et al.
even been found for mammals (Jarman 1974; Rob- 2004).
inson and Redford 1986; Rosenberger 1992) and
vertebrate predators (Marti 1993). Rare versus common species
In contrast, the humped-shaped form of the
b-diversity proportion on the ecoregion level did Our second hypothesis was that the differences
not support our expected linear increase with body between functional guilds should increase with
size, which suggests that the relationship between increasing weight placed on more-abundant species.
body size and host specificity might be weak for We consistently found that higher spatial scales were
saproxylic beetles. Thus, body size might not be a relatively more important for species turnover of
good predictor of geographical range for some insect abundant species, whereas the turnover of rare
taxa (Blackburn and Gaston 2003; Chown and Gaston species occurred more on small spatial scales. These
2010), which is in line with the results of Summer- results suggest that the availability of suitable hab-
ville et al. (2006), who found no effects of body size itats for rare species on small spatial scales varies
(small vs. large) on the diversity partitioning of forest greatly, which leads to a species turnover of the same
moths in deciduous forest of North America. Other magnitude as among ecoregions. If this result holds
taxa also do not show a positive correlation between true even in future studies that include other highly
body size and host range, e.g., non-predatory hover- diverse taxa, it would be highly relevant for conser-
flies (Gilbert 1990), tephrid flies (Kubota et al. 2007), vation because it emphasizes the importance of
newts (Joly and Giacoma 1992; Braz and Joly 1994), structural diversity on small spatial scales. In con-
and birds (Brandl et al. 1994), which could have trast, for more-abundant species, small spatial scales
several explanations. The main difference between are less important. At this point, we also have to keep
the results of our two approaches (body-size vs. host in mind the influence of our choice of methodology.
specificity) for studying the b-diversity of saproxylic An application of an additive approach may influence
beetles was that the influence of the ecoregion scale the conclusions here. Additive approaches will gen-
decreased as the host specificity decreased, whereas erally reveal higher importance of the ecoregional
the influence of the ecoregion scale was higher for level (Fig. S4-1; S4-2) because multiplicative parti-
species of intermediate body size than for species of tions are sensitive only to joint species presences and
small and large body size. These inconsistent patterns ignore joint absences; in contrast, additive partitions
could be caused by differences in the life-history are influenced by both. This is an important differ-
traits of the different beetle species. Some small ence in the property of multiplicative and additive
species might be highly vagile, fecund, and generalist partitions (Tuomisto 2010).
feeders, e.g., some bark beetles (Ranius 2006), which
would lead to a wide distributional range. Some large Implications for nature conservation
species might depend on specific resources or struc-
tures, such as rot holes, e.g., Osmoderma eremita For conservation strategies, we can conclude from
(Ranius and Heding 2001), which they occupy for our results that the influence of spatial scales on
many years and thus exhibit less willingness to species turnover is clearly trait dependent and
disperse. Moreover, the body size of saproxylic depends on whether more weight is given to rare or
beetles is highly constrained by phylogeny, and this more-common species. Several authors have recently
123Landscape Ecol (2011) 26:411–424 421
stressed that conservationists should focus more on bugs and saproxylic beetles was more strongly
common species to sustain ecosystem functions affected than species turnover of more generalist
because their loss rapidly alters ecosystems (e.g., species. This suggests that forest management should
Taylor et al. 2006). In contrast, all current efforts to ensure the sustainability of resources that are unique
conserve, e.g., saproxylic beetles, are focused only on within an ecoregion and therefore important for a
a handful of rare, mostly large and conspicuous specialized insect community because these species
species. For example, the ecological network of often have even smaller home ranges than their host
protected areas in the European Union (Natura 2000) plants or distribution of habitat structures. (5) Con-
considers only eight (of more than 1,300) rare servation strategies that only focus on a few con-
saproxylic beetle species as target species for spicuous and rare species do not consider the
conservation in Germany; seven of these are large important scales of whole species compositions and
(10–50 mm), one is medium sized (5 mm), and none may fail to save important ecosystem functions and
are small sized (\2.5 mm) (Petersen et al. 2003). processes.
This indicates that the current conservation efforts
does not consider the natural variability in species Acknowledgments We are grateful to all the researchers
who contributed to the projects from which data was compiled
traits and also neglects the importance of common for the present study. We thank the Bavarian State Institute of
species for biodiversity functions (Gaston 2010). The Forestry for providing data, Karen A. Brune for linguistic
necessity of considering these trait-based groups in revision of the manuscript, and two anonymous referees for
conservation is supported by a few other diversity- their critical comments and helpful suggestions on a previous
draft.
partitioning studies that used different measures of
diversity with different weights for rare and abundant
species (Gering et al. 2003; Summerville et al. 2003;
Müller and Gossner 2010). References
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