Received: 26 May 2021 | Revised: 3 November 2021 | Accepted: 8 November 2021

DOI: 10.1002/ece3.8393

RESEARCH ARTICLE

Influence of tree hollow characteristics and forest structure on

saproxylic beetle diversity in tree hollows in managed forests
in a regional comparison

Benjamin Henneberg1,2 | Simon Bauer2 | Markus Birkenbach2 | Vanilla Mertl2 |

Manuel J. Steinbauer3 | Heike Feldhaar1 | Elisabeth Obermaier2

1
Department of Animal Ecology I, Bayreuth
Center of Ecology and Environmental Abstract
Research (BayCEER), University of Bayreuth, Tree hollows are among the rarest habitats in today's Central European managed for-
Bayreuth, Germany
2 ests but are considered key structures for high biodiversity in forests. To analyze and
Ecological-­Botanical Garden of the
University of Bayreuth, Bayreuth, Germany compare the effects of tree hollow characteristics and forest structure on diversity
3
Department of Sport Ecology, Bayreuth of saproxylic beetles in tree hollows in differently structured managed forests, we
Center of Ecology and Environmental
Research (BayCEER), University of Bayreuth,
examined between 41 and 50 tree hollows in beech trees in each of three state forest
Bayreuth, Germany management districts in Germany. During the two-­year study, we collected 283 sap-

Correspondence
roxylic beetle species (5880 individuals; 22% threatened species), using emergence
Benjamin Henneberg, Department of Animal traps. At small spatial scales, the size of hollow entrance and the number of surround-
Ecology I, Bayreuth Center of Ecology
and Environmental Research (BayCEER),
ing microhabitat structures positively influenced beetle diversity, while the stage of
University of Bayreuth, Universitätsstraße wood mould decomposition had a negative influence, across all three forest districts.
30, 95444 Bayreuth, Germany.
Email: benjamin1.henneberg@uni-bayreuth.
We utilized forest inventory data to analyze the effects of forest structure in radii
de of 50–­500 m around tree hollows on saproxylic beetle diversity in the hollows. At

Funding information
these larger spatial scales, the three forest management districts differed remark-
Landesanstalt für Wald-­und Forstwirtschaft ably regarding the parameters that influenced saproxylic beetle diversity in tree hol-
(LWF), Grant/Award Number: L058;
Oberfrankenstiftung; German Research
lows. In Ebrach, characterized by mostly deciduous trees, the amount of dead wood
Foundation; University of Bayreuth positively influenced beetle diversity. In the mostly coniferous Fichtelberg forest
district, with highly isolated tree hollows, in contrast, only the proportion of beech
trees around the focal tree hollows showed a positive influence on beetle diversity.
In Kelheim, characterized by mixed forest stands, there were no significant relation-
ships between forest structure and beetle diversity in tree hollows. In this study, the
same local tree hollow parameters influenced saproxylic beetle diversity in all three
study regions, while parameters of forest structure at larger spatial scales differed in
their importance, depending on tree-­species composition.

KEYWORDS

Coleoptera, conservation, dead wood, threatened species, tree hollows, wood mould

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,
provided the original work is properly cited.
© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

Ecology and Evolution. 2021;11:17973–17999.  www.ecolevol.org | 17973

17974 | HENNEBERG et al.

1 | I NTRO D U C TI O N (Micó, 2018; Micó et al., 2015; Quinto et al., 2014; Schauer et al.,
2018). Besides limited habitat availability, many saproxylic beetles
Recent studies have shown that the abundance and species richness are assumed to have limited dispersal abilities (Feldhaar & Schauer,
of insects have declined strongly over past decades (Wagner, 2020). 2018). Therefore, it is important to analyze the influence of forest
This is reported for agricultural landscapes (Wagner, 2020) as well parameters on saproxylic beetle diversity in tree hollows at different
as managed forest ecosystems (Seibold et al., 2019). Intensive for- spatial scales (Ranius et al., 2015), from characteristics of the tree
est management has resulted in a massive decline of key elements hollows themselves (Quinto et al., 2014; Schauer et al., 2018), to pa-
important for forest biodiversity, such as dead or moribund trees rameters of forest structure such as distribution of dead wood, age
(Vogel et al., 2020). Dead wood is a characteristic and abundant re- structure, and tree-­species composition (Floren et al., 2014; Gossner
source in natural forest ecosystems (Seibold & Thorn, 2018) but is et al., 2013; Micó et al., 2013).
often removed in European managed forests (Thorn et al., 2020), The aim of this study was to assess the influence of parameters
resulting in low amounts of dead wood (Gossner, Lachat, et al., 2013; of tree hollows and the surrounding forest structure on the diversity
Thorn et al., 2020). Reducing the amount of dead wood in managed of saproxylic beetles in tree hollows at different spatial scales. We
forests can help lowering the risk of natural disturbances like wild- conducted our study in a regional comparison to achieve general-
fires through fuel reduction or pest insect outbreaks (Leverkus et al., izable results and to investigate possible regional differences. We
2020). However, this practice also eliminates important habitat fea- selected three state forest regions in Bavaria, Germany, that differed
tures for saproxylic taxa (Thorn et al., 2016). Consequently, many in tree-­species composition but were all representative for Central
species of saproxylic insects, which directly or indirectly depend on European managed forests. In addition to local parameters of tree
dead wood in at least one stage of their life cycle (Speight, 1989), hollows and surrounding forest structure we recorded in the field,
have become endangered or extinct (Seibold et al., 2015; Thorn we used forest inventory data that are collected systematically for
et al., 2020). Since approximately 34% of forest-­dwelling species in all state forest regions by the Bavarian state forest authority (BaySF)
Europe are regarded as saproxylic (Müller et al., 2008), this also con- and other German state forest authorities. The use of forest inven-
tributes to the decline of forest biodiversity in general. Beetles are tory data allowed us to statistically analyze parameters of forest
especially threatened as >50% of all forest-­dwelling beetle species in structure at larger spatial scales around the focal tree hollows.
Germany are regarded as saproxylic (Köhler, 2000). Here we address the following questions: (I) Which parameters
Large old trees containing tree hollows are among the rarest of tree hollows and forest structure of managed forests are related
structures in European managed forests (Lindenmayer et al., 2012). to species richness of saproxylic beetles in tree hollows of beech
Tree age and diameter are important factors that facilitate the devel- trees at different spatial scales? Based on existing studies we hy-
opment of tree hollows (Ranius et al., 2009). In contrast to other dead pothesize that local tree hollow parameters like area of hollow en-
wood structures like logs, tree hollows provide long-­lasting microhab- trance (Quinto et al., 2014) or temperature inside the hollows (Müller
itats that offer nutritional resources for many species of saproxylic et al., 2015; Schauer et al., 2018) as well as early to intermediate
organisms (Siitonen, 2012), and a specific abiotic environment charac- stages of wood mould decomposition (Schauer et al., 2018) will posi-
terized by stable temperature and moisture conditions and increased tively influence saproxylic beetle species richness, while the amount
pH values (Müller et al., 2014). Each tree hollow is unique with regard of dead wood might positively influence beetle species richness at
to the combination of microenvironmental characteristics and the larger spatial scales (Müller et al., 2015). (II) Are there common pa-
range of microhabitats within (Quinto et al., 2014; Siitonen, 2012). rameters explaining species richness of hollow-­using beetles across
Besides dead wood generalists, tree hollows also provide habitat for all three forest regions? We hypothesize that local tree hollow pa-
many highly specialized saproxylic beetle species (Speight, 1989). rameters might be more influential to saproxylic beetle species rich-
These cavity-­dependent tree hollow specialist species complete ness than parameters of forest structure at larger spatial scales as
most of their life cycle in tree hollows and represent an exceptionally the quality of tree hollows is crucial for the development of beetle
threatened group with approximately 75% of them being considered individuals. Tree hollows provide nutritional resources as well as a
to be threatened or endangered (Schmidl & Büche, 2018). diversity of microhabitats suitable for the development of saproxylic
In past decades European forests were mainly managed for beetles while parameters of forest structure at larger spatial scales
timber production. Since the late 1990s, biodiversity conservation might be more important for accessibility of tree hollows and popu-
programs with a special focus on dead wood structures have been lation sizes of saproxylic beetles.
implemented in an increasing number of European countries, includ-
ing Finland, Sweden, and Germany (Thorn et al., 2020; Vítková et al.,
2018). However, to implement effective conservation measures 2 | M ATE R I A L S A N D M E TH O DS
it is crucial to better understand which forest parameters are the
most important to enhance saproxylic beetle diversity. While sev- 2.1 | Study sites
eral studies have analyzed the habitat requirements of saproxylic
beetles in general (Müller et al., 2015; Ranius et al., 2015; Seibold The study was conducted in 2018 and 2019 in three Bavarian state
et al., 2016), less is known about beetle communities in tree hollows forest management districts (Bayerische Staatsforsten, BaySF):
HENNEBERG et al. | 17975

Ebrach (N 49°50′, E 10°29′), Fichtelberg (N 49°59′, E 11°50′), and The black acrylic mesh also did not allow insects to pass. During the
Kelheim (N 48°55′, E 11°52′). These forest management districts sampling period from April to September (18 weeks), all tree hol-
were chosen because they represent typical managed forest types lows were closed with black fabric and emergence traps (modified
in Central Europe as they display the full range of management in- from Gouix & Brustel, 2012) (Figure A4) that allow efficient sampling
tensity from strict forest reserves to intensively managed forests of tree hollow arthropod communities as only individuals emerg-
(Gossner, Lachat, et al., 2013). The study regions also represent a ing from the tree hollows will be trapped (Schauer et al., 2018). The
gradient in tree-­species composition from semi-­natural beech for- collecting bottles contained 99.8% ethanol and were emptied bi-
ests (Ebrach) to mixed forests (Kelheim) and forests with a high pro- weekly. A beetle taxonomist (Boris Büche) identified all beetles to
portion of planted Picea abies trees (Fichtelberg) that is typical for species-­level.
Central European managed forests (Müller et al., 2008).
The forest management district Ebrach in northern Bavaria
consists of temperate deciduous forest stands (app. 1000 km2, low 2.3 | Parameters of tree hollows/hollow-­bearing
mountain range, mean annual temperature: 7–­8°C, mean annual pre- trees recorded in the field
cipitation: 850 mm [Bässler et al., 2014]). The dominant tree species
is beech Fagus sylvatica (43% cover), followed by oak (Quercus robur The following parameters of each tree hollow/hollow-­bearing tree
and Quercus petraea, 20%). Deciduous trees cover more than 70% were recorded:
of the forest area (Müller et al., 2008). The altitude of sample trees
ranged from 324 to 482 m a.s.l. • Area of hollow entrance calculated as the area of an ellipse with
The forest management district Fichtelberg, located in the low A = π*a*b, where a is half the height and b half the width of the
mountain range Fichtelgebirge, consists of mainly coniferous for- hollow entrance.
est stands (app. 157 km2, mean annual temperature: 5–­6°C, mean • Hollow volume calculated as the volume of a cylinder with
annual precipitation: 1000–­1500 mm [BaySF, 2017]), and is charac- V = π*r2*h, where r is the internal radius of the hollow measured
terized by humid, sub-­alpine climate. The dominant tree species is at the entrance and h the internal height of the hollow measured
spruce P. abies (80% cover), followed by beech (7%) (BaySF, 2017). with a telescopic measuring stick.
The altitude of sample trees ranged from 525 to 873 m a.s.l. • Height above ground measured as distance of the lowest point of
The forest management district Kelheim consists of mixed for- the hollow entrance to the ground.
est stands (app. 179 km2, mean annual temperature: 7–­8°C, mean • Diameter at breast height (DBH) of the hollow tree measured at
annual precipitation: 650–­850 mm [BaySF, 2015]), and is charac- 130 cm above ground.
terized by sub-­oceanic climate. Its forest stands are mixed in tree-­ • Height above sea level: altitude of each tree hollow in m a.s.l.
species composition with 56% coniferous and 44% deciduous trees. • Stage of decomposition: the stage of decomposition of wood mould
The dominant tree species is spruce (44% cover), followed by beech sampled from the base of each tree hollow at a depth of 2–­5 cm,
(29%) (BaySF, 2015). The altitude of sample trees ranged from 396 using a spoon that was attached to a stick, was determined using
to 566 m a.s.l. three parameters, color, texture, and visible woody parts, and
We selected between 41 and 50 beech trees with tree hollows was classified in four ascending categories according to Jarzabek
in each forest management district (Ebrach: 50, Fichtelberg: 43, (2005):
Kelheim: 41) that were distributed over the whole area of each man- Stage 1/low decay: yellow to light brown in color, visible woody
agement district (Figures A1–­A3). Tree hollows were selected if they parts of bigger size.
contained at least 2 cm of wood mould at the bottom of the hollow, Stage 2/medium decay: light brown to brown in color, visible
and the diameter at breast height (DBH) of the host tree was at least woody parts of smaller size.
20 cm. Only tree hollows with a maximum height above ground of Stage 3/medium to high decay: brown to dark brown in color, al-
the lowest point of the hollow entrance of 350 cm were sampled. most no visible woody parts.
The minimum distance between two sample trees was 200 m, and Stage 4/high decay: dark brown to black in color, no visible woody
the minimum distance to the forest edge was 100 m. We randomly parts.
selected tree hollows matching the criteria in each forest stand by as- • Temperature inside the hollow measured with temperature loggers
signing each tree hollow in a given stand a number and rolling a dice. inside each tree hollow every 60 min over the whole sampling
period.

2.2 | Sampling method and identification of

saproxylic beetles 2.4 | Parameters of forest structure recorded
in the field
After selection in February and March (Ebrach/Fichtelberg: 2018,
Kelheim: 2019), all tree hollows were closed with black acrylic mesh Two parameters of forest structure were recorded around each tree
to prevent vertebrates like birds from using them as nesting place. hollow:
17976 | HENNEBERG et al.

• Surrounding microhabitat structures: the number of microhabitat 2.7 | Statistical analysis of forest inventory data
structures in trees according to Kraus et al. (2016) in a 30 m radius
around each tree hollow: woodpecker holes, visible tree fungi, Interpolation maps displaying forest inventory parameters cannot
broken branches with a minimum diameter of 12 cm, and injuries be directly used for statistical analysis as they display a spatially
to the bark with a minimum area of 250 cm2. These tree-­related continuous data distribution. To obtain values for each sampled
microhabitat structures have been widely recognized as import- tree hollow we used the software ArcGIS (ESRI, 2018) to transform
ant substrates and structures for saproxylic biodiversity in forests the interpolated data to a point grid of 10 × 10 m. We drew circu-
and are receiving increasing attention in management, conserva- lar buffers around each tree hollow with radii ranging from 50 to
tion and research (Larrieu et al., 2018). 500 m and calculated the average value for each forest inventory
• Surrounding tree hollows: the number of visible tree hollows in a parameter within each buffer. We used these values for the statis-
30 m radius around each tree hollow. tical analysis to examine the forest inventory parameters at differ-
ent spatial scales around each tree hollow. We randomly excluded
buffer sizes or single tree hollows from the analysis when there was
2.5 | Parameters of forest structure assessed via more than 10% overlap of the buffer areas of neighboring tree hol-
forest inventory data lows or with the border of the forest management district.

We used forest inventory data collected by the Bavarian state for-

est authorities (BaySF) in 2010–­2012. Sampling of forest inventory 2.8 | Statistical analysis
data is conducted every 10 years by the BaySF on a 200 × 200 m
grid over the whole area of the forest management district, with To visualize similarities in species composition between the three
a sampling point at each nodal point. More than 100 parameters study sites, detrended correspondence analysis (DCA) was per-
are recorded within a parameter-­d ependent radius around each formed using the R function decorana in the vegan package (Oksanen
sampling point, out of which we chose those that we expected to et al., 2013). To analyze the influence of tree hollow parameters
influence saproxylic beetle diversity: amount of dead wood (vol- and parameters of forest structure on species richness of hollow-­
ume of all dead woody parts, standing or downed, with a minimum using beetles, generalized linear models (GLMs) with Poisson error
2
diameter of 20 cm), sampled within a radius of 12.62 m (500 m ) distribution were implemented. Collinearity among explanatory
around each sampling point, and DBH of deciduous trees (the mean variables was tested using Pearson's linear correlation (exclusion
diameter at breast height in cm) as a proxy for age structure of criterion for two variables in the same model |r| > .7). Univariate
deciduous trees, also sampled within a radius of 12.62 m (500 m2) GLMs were used for preselection of explanatory variables for mul-
around each sampling point. Additionally, in the forest manage- tivariate GLMs. All explanatory variables with significant influence
ment district Fichtelberg that consisted mainly of coniferous trees, on species richness in univariate GLMs were used in combination
we examined the proportion of beech trees. For parameters DBH of with forest inventory data in multivariate GLMs. If two variables
deciduous trees and proportion of beech trees, only trees assigned to showed collinearity they were used in separate models. If there
the upper forest layer were included in the analysis to avoid over- were no local tree hollow parameters with significant influence on
representation of young trees that lack suitable microhabitats for species richness in univariate GLMs, the four parameters with the
saproxylic beetles. lowest p-­values were selected for multivariate GLMs. To improve
model fitting and the distribution of residuals all explanatory vari-
ables were transformed. Depending on the data distribution, either
2.6 | Interpolation of forest inventory data a log-­or sqrt-­transformation was chosen. By visually inspecting the
distribution of residuals in univariate GLMs the model fit of trans-
Because forest inventory data is recorded as point data, it must be formed variables was compared to the untransformed variables,
interpolated to maps with a continuous spatial distribution prior to and the version that provided the best distribution of residuals was
regression analysis with species richness of saproxylic beetles. We selected for multivariate GLMs. Stepwise model selection based on
tested the forest inventory parameters for spatial autocorrelation AIC was used to remove non-­relevant variables from the models
by visually inspecting semivariograms in the software ArcGIS (ESRI, (R function step in the stats package). All multivariate models were
2018). The only parameter of forest inventory data where we de- tested for overdispersion using the R function dispersion.test in the
tected spatial autocorrelation and that also had a normal distribu- AER package (Kleiber & Zeileis, 2008). Each explanatory variable
tion, and we therefore were able to apply the geostatistical Kriging that showed significant influence on species richness in multivari-
approach was DBH of deciduous trees in the forest management dis- ate GLMs was visualized using the R package visreg (Breheny &
trict Ebrach. For all other forest inventory parameters and forest Burchett, 2017). Analyses were performed for total species rich-
management districts we applied the deterministic inverse distance ness of saproxylic beetles as well as for the subset of threatened
weighting (IDW) method to spatially interpolate forest inventory species (i.e. species listed with a status of 0–­3 on the Red List of
data and create interpolation maps (Figures A5–­A6). Germany (Schmidl & Büche, 2018)).
HENNEBERG et al. | 17977

To combine all three forest management districts in a single

model we created generalized linear mixed-­effects models (GLME)
using the R function glmer in the lme4 package (Bates et al., 2015),
with parameters of tree hollows and forest structure as fixed
factors and forest management district as the random factor.
Residuals of random factors were examined visually by inspecting
the fitted vs. residuals plot (R function plot in the lme4 package
[Bates et al., 2015]) and the residuals vs. predicted plot (R function
simulateResiduals in the DHARMa package [Hartig, 2020]). All anal-
yses were performed with the software R, version 3.6.3 (R Core
Team, 2020).

F I G U R E 1 Detrended correspondence analysis (DCA) that

3 | R E S U LT S visualizes similarities in species composition between the three
study sites; circles = tree hollows, crosses = saproxylic beetle
3.1 | Saproxylic beetles species

In the three forest management districts (134 tree hollows), we

collected a total of 5880 saproxylic beetle individuals, belong- p < .05), and to the stage of decomposition of the wood mould inside
ing to 283 species and 48 families. Sixty-­t wo species (21.9%) are the hollows (z = −2.82, p < .001). There was a hump-­shaped rela-
regarded as threatened (Schmidl & Büche, 2018). Four species of tionship to temperature inside the hollows (z = 3.82, p < .001). This
the collected beetle species are critically endangered (Red List relationship turned positive in the r = 500 m model. Furthermore,
of Germany status 1 [Schmidl & Büche, 2018]): Ampedus brunni- representing forest structure at larger spatial scales, the number of
cornis, Crepidophorus mutilatus, Cryptophagus deubeli, Prionychus microhabitat structures in a 30 m radius around the tree hollows (sur-
melanarius. The sampled species are very diverse regarding body rounding microhabitat structures, z = 2.32, p < .05), and the amounts
size (ranging from 1–­2 mm to approximately 5 cm) and ecological of dead wood up to a radius of 100 m around the tree hollows (dead
guild (Schmidl & Bußler, 2004). Only few species are flightless; the wood volume, r = 50 m, z = 3.24, p < .01; r = 100 m, z = 2.72, p < .01)
majority is mobile. However, for most species it is not known if were positively correlated with the total number of saproxylic beetle
they are highly mobile or more or less sessile, and how far they species in the tree hollows (Figure 2, Table A1). Pseudo-­r2 values
disperse within a forest or even between forest regions (Feldhaar that show the explanatory power of the models range from 0.378 to
& Schauer, 2018). 0.740 depending on radius (Table A1).
In 2018 we collected 4151 individuals in Ebrach (50 tree hol- When only the 41 threatened saproxylic beetle species were
lows), belonging to 196 species from 43 families with 41 species included in the analysis, the volume of the tree hollows (z = 3.21,
(20.9%) being regarded as threatened. The same year we collected p < .01) was the only local tree hollow parameter that was pos-
441 individuals in Fichtelberg (43 tree hollows), belonging to itively correlated with the number of beetle species. The height
74 species from 24 families with nine species (12.2%) being re- of the hollow entrance above ground (z = −2.60, p < .01) and the
garded as threatened. In 2019 we collected 1288 individuals in stage of decomposition of the wood mould (z = −2.70, p < .01) were
Kelheim (41 tree hollows), belonging to 107 species from 32 fami- negatively correlated with the number of species. The number of
lies with 28 species (26.2%) being regarded as threatened. The av- microhabitat structures in a 30 m radius around the tree hollows
erage species richness of saproxylic beetle species per tree hollow (surrounding microhabitat structures, r = 30 m, z = 3.24, p < .01)
was 13.2 ± 6.0 (mean ± SD) in Ebrach, 4.6 ± 2.7 in Fichtelberg, and was the only parameter of forest structure that was positively cor-
7.9 ± 3.7 in Kelheim. Similarities in species composition between related with the number of threatened saproxylic beetle species
the three study sites were visualized using detrended correspon- (Figure A7, Table A2). Pseudo-­r 2 values that show the explanatory
dence analysis (DCA) (Figure 1). power of the models range from 0.288 to 0.691 depending on the
radius (Table A2).

3.2 | Effects of tree hollow characteristics and

forest structure on saproxylic beetle diversity 3.2.2 | Forest management district Fichtelberg

3.2.1 | Forest management district Ebrach In Fichtelberg species richness of saproxylic beetles was positively re-
lated only to the proportion of beech trees in the tree-­species compo-
In Ebrach species richness of saproxylic beetles was negatively sition at different spatial scales around the tree hollows (proportion of
related to the height of the tree hollows above ground (z = −2.31, beech trees, r = 50 m, z = 2.27, p < .05; r = 100 m, z = 2.28, p < .05)
17978 | HENNEBERG et al.

(Figure 3, Table A3). Thus, with increasing proportion of beech trees in 100 m around the focal tree hollows (r = 50 m, z = 2.54, p < .05;
the surrounding of tree hollows, the total number of hollow-­using bee- r = 100 m, z = 2.50, p < .05) (Figure A8, Table A4). Pseudo-­r2 values
2
tle species increased in this conifer dominated forest area. Pseudo-­r that show the explanatory power of the models range from 0.203 to
values that show the explanatory power of the models are 0.113 0.487 depending on radius (Table A4).
(r = 50 m) and 0.116 (r = 100 m) (Table A3).
When only the nine threatened saproxylic beetle species were
included in the analysis, species richness was positively related to 3.2.3 | Forest management district Kelheim
the number of microhabitat structures in a 30 m radius around the
tree hollows (surrounding microhabitat structures, r = 30 m, z = 2.19, In Kelheim species richness of saproxylic beetles showed a U-­
p < .05) and again the proportion of beech trees up to a radius of shaped relationship to the size of the area of hollow entrance

F I G U R E 2 Multivariate GLMs of the Ebrach management district (2018) of different radii around the focal tree hollows (r ≤ 100 m:
n = 50, r = 200 m: n = 37, r = 300 m: n = 32, r = 500 m: n = 24). Total species richness of saproxylic beetles as dependent variable; p < .05 (*),
p < .01 (**), p < .001 (***)
HENNEBERG et al. | 17979

temperature inside the hollows (z = 2.28, p < .05) (Figure A9, Table
A6). As there were no statistically significant parameters of forest
structure, Pseudo-­r2 that shows the explanatory power of the mod-
els is 0.293 for all radii (Table A6).

3.2.4 | Combined analysis of the three forest

management districts

Species richness of saproxylic beetles over all three management

districts combined was negatively related to local tree hollow
parameters height above sea level (z = −4.45, p < .001), height of
the hollow entrance above ground (z = −2.15, p < .05), and the
stage of decomposition of the wood mould (z = −2.19, p < .05), and
positively related to the area of hollow entrance (z = 4.17, p < .001).
The only parameter of forest structure that explained total species
richness across all three management districts was the number of
microhabitat structures in a 30 m radius around the tree hollows
(surrounding microhabitat structures, z = 2.70, p < .01) (Figure 5,
Table A7).
When only the 61 threatened saproxylic beetle species were in-
cluded in the analysis, species richness was positively related to the
volume of the tree hollows (z = 2.18, p < .05), and the temperature in-
side the hollows (z = 5.37, p < .001), and negatively related to height
above sea level (z = −5.47, p < .001). The only parameter describing
forest structure that explained species richness of threatened sap-
roxylic beetles across all three management districts was again the
number of microhabitat structures in a 30 m radius around the tree
hollows (surrounding microhabitat structures, z = 2.01, p < .05) (Figure
A10, Table A8).

F I G U R E 3 Multivariate GLMs of the Fichtelberg management

district (2018) of different radii around the focal tree hollows 4 | D I S CU S S I O N
(r ≤ 100 m: n = 43, r = 200 m: n = 36, r = 300 m: n = 33, r = 500 m:
n = 28). Total species richness of saproxylic beetles as the In this study, the influence of local tree hollow parameters as well
dependent variable; p < .05 (*), p < .01 (**), p < .001 (***)
as the surrounding forest structure within three managed forests
on species richness of saproxylic beetles in tree hollows was in-
vestigated. Our results confirm the findings that species richness
(z = −2.23, p < .05). This can be explained by a single tree hol- of hollow-­using saproxylic beetles is related on the one hand to
low with a small entrance area. When that hollow was removed different properties of the tree hollows themselves (Micó et al.,
from the analyses, the relationship became positive. There was a 2015; Quinto et al., 2014; Schauer et al., 2018): when all three
negative relationship between species richness and height of the forest regions were combined the area of hollow entrance posi-
hollow entrance above ground (z = −1.96, p < .05). The number of tively influenced total species richness while height above ground
microhabitat structures in a 30 m radius around the tree hollows and stage of decomposition negatively influenced total species
(surrounding microhabitat structures, r = 30 m, z = −3.16, p < .01) richness of saproxylic beetles; on the other hand, the influence of
also showed a U-­shaped relationship to total species richness these local properties of tree hollows on species richness of sap-
(Figure 4, Table A5). Pseudo-­r 2 values that show the explanatory roxylic beetles also depends on surrounding forest structure. In
power of the models range from 0.240 to 0.434 depending on the forest regions with a higher proportion of deciduous trees (Ebrach,
radius (Table A5). Kelheim) the influence of local tree hollow parameters was more
Species richness of 28 threatened saproxylic beetle species pronounced than in the mostly coniferous Fichtelberg forest re-
was positively related to the volume of the tree hollows (z = 3.21, gion. When regions were analyzed separately, the only parameter
p < .01), the area of hollow entrance (z = 2.54, p < .05) and the (of those investigated) that influenced total species richness of
17980 | HENNEBERG et al.

F I G U R E 4 Multivariate GLMs of the Kelheim management district (2019) of different radii around the focal tree hollows (r ≤ 100 m:
n = 41, r = 200 m: n = 35, r = 300 m: n = 32). Total species richness of saproxylic beetles as the dependent variable; p < .05 (*), p < .01 (**),
p < .001 (***)

saproxylic beetles in the conifer-­d ominated Fichtelberg region 4.1 | Influence of local tree hollow parameters
was the proportion of beech trees in the surrounding of focal tree
hollows. Beech trees in this forest region can be described as iso- One local tree hollow parameter that showed a positive relation-
lated within a matrix of coniferous trees which made their pres- ship to species richness when all three forest regions were com-
ence the most important resource for saproxylic beetles under bined was area of hollow entrance. A positive relationship with
these circumstances. Therefore, the influence of local tree hollow area of hollow entrance was also observed for threatened species
characteristics on diversity of saproxylic beetles must be seen in in Kelheim. This relationship has also been observed in previous
relation to surrounding forest structure. However, the sample size studies (Quinto et al., 2014; Ranius, 2002; Schauer et al., 2018).
is potentially limited for the analysis of each region separately, It has been proposed that many saproxylic species prefer a less
as there were only 41–­5 0 tree hollows examined in each forest humid microclimateand a larger hollow entrance is associated with
region. a reduction in humidity inside the tree hollow (Schauer et al., 2018).
HENNEBERG et al. | 17981

F I G U R E 5 Generalized linear mixed

effects models (GLMEs) of the three
forest management districts combined
(n = 134 tree hollows). Total species
richness of saproxylic beetles as the
dependent variable; p < .05 (*), p < .01 (**),
p < .001 (***)

We also found a positive relationship between hollow volume also reported a positive relationship between temperature and spe-
and number of threatened species in all three forest regions com- cies richness (Müller et al., 2015; Schauer et al., 2018), especially for
bined, as well as in Ebrach and Kelheim alone. As many threatened threatened saproxylic beetle species (Lindhe et al., 2005; Widerberg
saproxylic beetle species are assumed to have high habitat re- et al., 2012).
quirements concerning not only quantity but quality of dead wood In Ebrach and Kelheim we found higher species richness in tree
structures (Müller et al., 2005), several studies have reported the hollows located closer to the ground. We observed the same rela-
association of threatened species with large tree hollows that offer a tionship for total species richness in all three forest regions com-
greater diversity of microhabitats (Ranius et al., 2009). bined as well as for threatened species in Ebrach. This has also
There was a negative relationship between total species richness been reported by Schauer et al. (2018) for the Ebrach management
and the stage of wood mould decomposition when all three manage- district as well as Quinto et al. (2014) in Mediterranean deciduous
ment districts were combined as well as for Ebrach alone for all spe- oak woodland. One possible explanation could be the higher num-
cies as well as for threatened species. A similar relationship has been ber of generalist predator species in tree hollows located closer to
reported by Stokland and Siitonen (2012), who proposed that the the ground (Ranius, 2002; Ranius, Svensson, et al., 2009). It has also
number of saproxylic species in forest dead wood declined at later been reported that generalist species that can dwell on the forest
stages of decay. While we found early stages of wood mould de- floor are more dominant in tree hollows that are connected to the
composition to contain the highest species richness, both Sverdrup-­ ground (Siitonen, 2012). However, as we only examined tree hollows
Thygeson et al. (2010) and Schauer et al. (2018) reported the highest up to a height of 350 cm above ground, we only cover a limited ver-
species richness in tree hollows with intermediate stages of wood tical range of tree hollows, which may bias results.
mould decomposition. Total and threatened species richness for all three manage-
In Ebrach total species richness of saproxylic beetles was related ment districts combined was negatively related to height of the
to temperature inside the tree hollows. We found the highest spe- tree hollows above sea level. It has previously been shown that
cies richness in tree hollows with intermediate mean temperatures altitude negatively affected saproxylic beetle diversity (Johansson
of 17–­18°C. The positive relationship between temperature and et al., 2017; Wu et al., 2008) as higher elevations are generally
species richness in the r = 500 m model can be explained by three associated with colder climate. Johansson et al. (2017) described
hollows with a relatively high average temperature that contained gradients in altitude as important determinants of the distribu-
relatively few beetle species and that were randomly removed from tion of saproxylic beetle assemblages in boreal forests in Sweden.
the r = 500 m model due to spatial overlap with other hollows or Similarly, Müller et al. (2015) proposed that conservationists, land-
non-­forest areas. However, for threatened species in all three for- scape managers, and ecologists should pay more attention to the
est regions combined, as well as in Kelheim alone, we observed a climate gradient as one fundamental driver of saproxylic insect
strictly positive relationship with temperature. Many other studies diversity.
17982 | HENNEBERG et al.

4.2 | Influence of forest structure at larger most important parameter influencing saproxylic beetle diversity
spatial scales in tree hollows, and the quality of the habitat –­ the tree hollow
itself –­might be of secondary importance.
Beside parameters of the tree hollows themselves, we also ana- Implications for forest management that can be inferred
lyzed forest structure at different spatial scales. In contrast to from this study include the conservation of developing and ex-
tree hollow parameters, the parameters of forest structure that isting tree hollows as each tree hollow is unique with regard to
influenced saproxylic beetle diversity in tree hollows were mostly the range of microhabitats within (Quinto et al., 2014; Siitonen,
highly idiosyncratic with respect to region, which was most likely 2012). This study has also shown that the number of tree-­r elated
due to the differences in tree-­species composition between microhabitats (e. g. woodpecker holes, injuries to the bark) in
regions. a forest stand is not only beneficial to the development of fu-
It has previously been reported that the number of tree hollows ture tree hollows but seems to already increase species richness
in the surrounding of a focal tree hollow had a positive influence of saproxylic beetles in tree hollows and should therefore be
on species richness of saproxylic beetles (Ranius & Wilander, 2000; conserved. Another important implication for forest manage-
Schauer et al., 2018). In our study we could not confirm this, but the ment is the further enrichment of dead wood in managed for-
number of microhabitat structures, indicating the number of pos- ests (Gossner, Floren, et al., 2013; Müller et al., 2015), especially
sible future tree hollows or other dead wood related structures to dead wood from deciduous tree species as saproxylic beetles
develop, in a 30 m radius around each focal tree hollow explained have been shown to react sensitively not only to the amount
species richness of saproxylic beetles across all three forest regions but also the variety and distribution of dead wood (Sverdrup-­
as well as for Ebrach and Kelheim alone. The same result was ob- Thygeson et al., 2014). Tree-­s pecies composition has also proven
tained for threatened species in all three regions combined as well as to greatly influence diversity of hollow-­u sing beetles (Floren
for Ebrach and Fichtelberg alone. et al., 2014). Therefore, increasing the proportion of deciduous
In Ebrach, the region with the highest proportion of decidu- trees in managed forests with a high proportion of coniferous
ous tree species, species richness of saproxylic beetles showed trees will increase species richness of saproxylic beetles in tree
a significant positive relationship with dead wood volume at hollows.
small radii of 50 and 100 m around the focal tree hollows but
not at larger spatial scales. A reason for the restriction of the
influence of dead wood volume to smaller spatial scales might be 5 | CO N C LU S I O N S
that dispersal abilities of most saproxylic insect species are still
unknown and might be rather small (Feldhaar & Schauer, 2018). Local tree hollow characteristics influence the diversity of saprox-
In the other two forest management districts, with fewer beech ylic beetles in tree hollows irrespective of region and management.
trees and a lower proportion of deciduous trees, there was no Parameters of forest structure at larger spatial scales differ in their
relationship between dead wood volume and species richness. importance for saproxylic beetle species richness depending on the
The importance of dead wood for a diverse fauna of saproxylic forest region's tree-­species composition. Therefore, the influence
insects in forests has been extensively described before (Floren of local tree hollow parameters on species richness of saproxylic
et al., 2014; Gossner, Floren, et al., 2013; Müller et al., 2015; beetles should be considered in the context of surrounding forest
Similä et al., 2003). Lassauce et al. (2011) proposed that addi- structure at larger spatial scales. Forest inventory data can sup-
tional variables like the type of dead wood or stage of decom- port this process by providing necessary data without additional
position should be taken into account. Stokland and Siitonen field work. Thus, forest inventory data can be a powerful tool, in
(2012) state that the majority of saproxylic beetle species (89%) combination with local tree hollow parameters, to assess diversity
are specialized either on deciduous or on coniferous dead wood. of hollow-­using beetles in managed forests and help forest authori-
Dead wood of mostly coniferous trees, as it prevailed in the for- ties decide which conservation measures to apply in certain parts
est management district Fichtelberg, did not promote species of forest regions to effectively protect saproxylic beetles in tree
richness of saproxylic beetles in tree hollows in beech trees. In hollows.
contrast, the only parameter that influenced total species rich-
ness of saproxylic beetles in the conifer dominated Fichtelberg AC K N OW L E D G M E N T S
management district was the proportion of beech trees in the The authors thank the Bavarian State Ministry of Food, Agriculture
surrounding of focal tree hollows, up to a radius of 100 m around and Forestry, the Landesanstalt für Wald-­ und Forstwirtschaft
tree hollows. This finding implies that in a forest region like the (LWF) (Project-­no. L058), and the Oberfrankenstiftung for fund-
Fichtelberg management district where suitable tree hollow hab- ing this research project. They thank the Government of Upper
itats in deciduous trees are very limited the number of poten- Franconia, Lower Franconia and Lower Bavaria for granting us
tial habitat trees in the surrounding of a focal tree hollow is the permission to conduct this study in Bavarian state forestries. They
HENNEBERG et al. | 17983

also thank the Bavarian state forestries (BaySF) for providing us filea​dmin/user_uploa​d/01-­ueber_uns/05-­s tand​orte/FB_Ficht​elber​
g/Natur​schut​zkonz​ept_Forst​betri​eb_Ficht​elberg.pdf
with forest inventory data, and Ulrich Mergner (Ebrach), Winfried
Breheny, P., & Burchett, W. (2017). Visualization of regression models
Pfahler and Martin Hertel (Fichtelberg), Sabine Bichlmaier using visreg. R Journal, 9, 56.
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of tree hollows. The authors thank Boris Büche for identifying the Research Institute.
Feldhaar, H., & Schauer, B. (2018). Dispersal of saproxylic insects. In
saproxylic beetles. They thank two anonymous reviewers for giv-
M. D. Ulyshen (Ed.), Saproxylic insects: Diversity, ecology and conser-
ing valuable comments on an earlier version of this manuscript. vation (pp. 515–­546). Springer.
This publication was funded by the German Research Foundation Floren, A., Muller, T., Dittrich, M., Weiss, M., & Linsenmair, K. E. (2014).
(DFG) and the University of Bayreuth in the funding program Open The influence of tree species, stratum and forest management on
Access Publishing. beetle assemblages responding to deadwood enrichment. Forest
Ecology and Management, 323, 57–­6 4.
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C O N FL I C T O F I N T E R E S T (2013). Effect of dead wood enrichment in the canopy and on
The authors declare that there are no conflicts of interest. the forest floor on beetle guild composition. Forest Ecology and
Management, 302, 404–­413. https://doi.org/10.1016/j.foreco.​
2013.​0 3.​0 39
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Science, 15, 251–­261.
ece3.8393
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APPENDIX A

FIGURE A1 Examined tree hollows in the Ebrach forest management district (n = 50)
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FIGURE A2 Examined tree hollows in the Fichtelberg forest management district (n = 43)
17988 | HENNEBERG et al.

FIGURE A3 Examined tree hollows in the Kelheim forest management district (n = 41)
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FIGURE A4 Emergence trap (modified from Gouix & Brustel,

2012)
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F I G U R E A 5 Distribution of dead wood in the Ebrach forest management district; map generated from forest inventory data using the
deterministic inverse distance weighting (IDW) method
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F I G U R E A 6 Proportion of beech trees in the Fichtelberg forest management district; map generated from forest inventory data using
the deterministic inverse distance weighting (IDW) method
17992 | HENNEBERG et al.

F I G U R E A 7 Multivariate GLMs of the Ebrach management district (2018) of different radii around the focal tree hollows (r ≤ 100 m:
n = 50, r = 200 m: n = 37, r = 300 m: n = 32, r = 500 m: n = 24). Species richness of threatened saproxylic beetles as the dependent variable;
p < .05 (*), p < .01 (**), p < .001 (***)
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F I G U R E A 8 Multivariate GLMs of the Fichtelberg management district (2018) of different radii around the focal tree hollows (r ≤ 100 m:
n = 43, r = 200 m: n = 36, r = 300 m: n = 33, r = 500 m: n = 28). Species richness of threatened saproxylic beetles as the dependent variable;
p < .05 (*), p < .01 (**), p < .001 (***)
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F I G U R E A 9 Multivariate GLMs of the Kelheim management district (2019) of different radii around the focal tree hollows (r ≤ 100 m:
n = 41, r = 200 m: n = 35, r = 300 m: n = 32). Species richness of threatened saproxylic beetles as the dependent variable; p < .05 (*), p < .01
(**), p < .001 (***)

F I G U R E A 1 0 Generalized linear mixed effects models (GLMEs) of the three forest management districts combined (n = 134 tree
hollows). Species richness of threatened saproxylic beetles as the dependent variable; p < .05 (*), p < .01 (**), p < .001 (***)
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|

TA B L E A 1 Multivariate GLMs of the Ebrach management district (2018) of different radii around the focal tree hollows (r ≤ 100 m: n = 50, r = 200 m: n = 37, r = 300 m: n = 32, r = 500 m:
n = 24)

GLM/radius Height above ground Stage of decomposition Temperature Microhabitat structures Dead wood volume (FI) Pseudo-­R2

r=0m n. s. p = .006** z = −2.746 p = .001** z = 3.220 n. a. n. a. .378

r = 50 m p = .021* z = −2.313 p = .005** z = −2.824 p = .0001*** z = 3.821 n. s. p = .001** z = 3.238 .507
r = 100 m p = .019* z = −2.343 p = .014* z = −2.446 p = .0001*** z = 3.799 n. s. p = .007** z = 2.717 .480
r = 200 m n. s. p = .0003*** z = −3.584 p = .002** z = 3.162 p = .021* z = 2.317 n. s. .457
r = 300 m n. s. p = 9.86e−06*** z = −4.420 p = 2.91e−05*** z = 4.180 p = .033* z = 2.136 n. s. .595
r = 500 m p = .030* z = −2.165 p = .015* z = −2.435 p = .003** z = 3.003 n. s. n. s. .740

Note: Total species richness of saproxylic beetles as dependent variable; p < .05 (*), p < .01 (**), p < .001 (***).
Abbreviations: n. a., not available; n. s., not significant.

TA B L E A 2 Multivariate GLMs of the Ebrach management district (2018) of different radii around the focal tree hollows (r ≤ 100 m: n = 50, r = 200 m: n = 37, r = 300 m: n = 32, r = 500 m:
n = 24)

GLM/radius Hollow volume Height above ground Stage of decomposition Microhabitat structures Pseudo-­R2

r=0m p = .008** z = 2.648 p = .017* z = −2.383 p = .028* z = −2.198 n. a. .288

r = 50 m n. s. p = .003** z = −2.989 p = .048* z = −1.980 p = .001** z = 3.214 .438
r = 100 m n. s. p = .004** z = −2.918 n. s. p = .001** z = 3.248 .443
r = 200 m n. s. p = .015* z = −2.441 p = .005** z = −2.780 p = .011* z = 2.554 .386
r = 300 m p = .009** z = 2.577 p = .011* z = −2.541 n. s. p = .015* z = 2.428 .496
r = 500 m n. s. p = .014* z = −2.468 p = .026* z = −2.234 p = .019* z = 2.349 .691

Note: Species richness of threatened saproxylic beetles as dependent variable; p < .05 (*), p < .01 (**), p < .001 (***).
Abbreviations: n. a., not available; n. s., not significant.
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TA B L E A 3 Multivariate GLMs of the Fichtelberg management district (2018) of different radii around the focal tree hollows (r ≤ 100 m:
n = 43, r = 200 m: n = 36, r = 300 m: n = 33, r = 500 m: n = 28)

GLM/radius Proportion of beech trees (FI) Pseudo-­R2

r=0m n. a.
r = 50 m p = .023* z = 2.269 .113
r = 100 m p = .023* z = 2.280 .116
r = 200 m n. s.
r = 300 m n. s.

Note: Total species richness of saproxylic beetles as dependent variable; p < .05 (*), p < .01 (**), p < .001 (***).
Abbreviations: n. a., not available; n. s., not significant.

TA B L E A 4 Multivariate GLMs of the Fichtelberg management district (2018) of different radii around the focal tree hollows (r ≤ 100 m:
n = 43, r = 200 m: n = 36, r = 300 m: n = 33, r = 500 m: n = 28)

GLM/radius Height above sea level Microhabitat structures Proportion of beech trees (FI) Pseudo-­R2

r=0m p = .004** z = −2.889 n. a. n. a. .287

r = 50 m n. s. p = .028* z = 2.196 p = .011* z = 2.540 .487
r = 100 m n. s. p = .027* z = 2.211 p = .012* z = 2.501 .483
r = 200 m n. s. p = .021* z = 2.306 n. s. .304
r = 300 m n. s. n. s. n. s. .203

Note: Species richness of threatened saproxylic beetles as dependent variable; p < .05 (*), p < .01 (**), p < .001 (***).
Abbreviations: n. a., not available; n. s., not significant.

TA B L E A 5 Multivariate GLMs of the Kelheim management district (2019) of different radii around the focal tree hollows (r ≤ 100 m:
n = 41, r = 200 m: n = 35, r = 300 m: n = 32)

GLM/radius Area of hollow entrance Height above ground Microhabitat structures Pseudo-­R2

r=0m p = .034* z = −2.121 p = .042* z = −2.035 n. a. .240

r = 50 m p = .049* z = −1.968 n. s. p = .002** z = −3.155 .434
r = 100 m p = .035* z = −2.105 n. s. p = .002** z = −3.145 .427
r = 200 m p = .026* z = −2.225 p = .050* z = −1.963 p = .001** z = −3.190 .388
r = 300 m p = .024* z = −2.255 p = .047* z = −1.985 p = .0007*** z = −3.408 .424

Note: Total species richness of saproxylic beetles as dependent variable; p < .05 (*), p < .01 (**), p < .001 (***).
Abbreviations: n. a., not available; n. s., not significant.

TA B L E A 6 Multivariate GLMs of the Kelheim management district (2019) of different radii around the focal tree hollows (r ≤ 100 m:
n = 41, r = 200 m: n = 35, r = 300 m: n = 32)

GLM/radius Area of hollow entrance Hollow volume Temperature Pseudo-­R2

r=0m p = .011* z = 2.542 p = .001** z = 3.210 p = .022* z = 2.287 .293

r = 50 m p = .011* z = 2.542 p = .001** z = 3.210 p = .022* z = 2.287 .293
r = 100 m p = .011* z = 2.542 p = .001** z = 3.210 p = .022* z = 2.287 .293
r = 200 m p = .011* z = 2.542 p = .001** z = 3.210 p = .022* z = 2.287 .293
r = 300 m p = .011* z = 2.542 p = .001** z = 3.210 p = .022* z = 2.287 .293

Note: Species richness of threatened saproxylic beetles as dependent variable; p < .05 (*), p < .01 (**), p < .001 (***).
Abbreviations: n. a., not available; n. s., not significant.
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TA B L E A 7 Generalized linear mixed-­effects models (GLMEs) of the three forest management districts combined (n = 134 tree hollows)

Random factor: Random factor: Random factor:

Ebrach Fichtelberg Kelheim Height above sea level

GLME total spp. Estimate = 0.187 Estimate = −0.104 Estimate = −0.080 p = 8.41e−06*** z = −4.454

Note: Total species richness of saproxylic beetles as dependent variable; p < .05 (*), p < .01 (**), p < .001 (***).
Abbreviations: n. a., not available; n. s., not significant.

TA B L E A 8 Generalized linear mixed-­effects models (GLMEs) of the three forest management districts combined (n = 134 tree hollows)

Random factor: Ebrach Random factor: Fichtelberg Random factor: Kelheim

GLME threatened spp. Estimate = 2.741e−16 Estimate = −1.844e−16 Estimate = −8.970e−17

Note: Species richness of threatened saproxylic beetles as dependent variable; p < .05 (*), p < .01 (**), p < .001 (***).
Abbreviations: n. a., not available; n. s., not significant.
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Area of hollow entrance Height above ground Stage of decomposition Microhabitat structures

p = 3.08e−05*** z = 4.168 p = .032* z = −2.145 p = .028* z = −2.193 p = .007** z = 2.701

Height above sea level Hollow volume Temperature Microhabitat structures

p = 4.61e−08*** z = −5.466 p = .029* z = 2.180 p = 7.99e−08*** z = 5.367 p = .045* z = 2.007