Forest Ecology and Management 553 (2024) 121609

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Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

How do stand features shape deadwood diversity?

Leszek Bujoczek a, *, Małgorzata Bujoczek b, Stanisław Zięba a
a
University of Agriculture in Krakow, Faculty of Forestry, Department of Forest Resources Management, Al. 29 Listopada 46, Krakow 31-425, Poland
b
University of Agriculture in Krakow, Faculty of Forestry, Department of Forest Biodiversity, Al. 29 Listopada 46, Krakow 31-425, Poland

A R T I C L E I N F O A B S T R A C T

Keywords: Saproxylic species from different taxonomic groups often occur only on certain types of deadwood with specific
Deadwood diversity qualitative characteristics. The various types of deadwood are very dynamic elements of forest ecosystems,
Saproxylic organisms associated with many site and stand features, as well as with the type of forest management. Using a pool of
Coarse woody debris
29,098 sample plots spread across Poland, we analyzed 30 different deadwood types defined on the basis of three
Forests
Biodiversity protection
characteristics: position (standing, lying), degree of decomposition, and size. Statistical hurdle models were used
to assess changes in the volume of individual deadwood types based on a broad range of independent variables.
Depending on the type of management, terrain, site fertility, stand volume, tree density, and stand age, the
models revealed substantial differences in the volume of different deadwood types, ranging from 0 to approx. 4
m3 ha− 1. It was found that the volume of most deadwood types (except for a few, mostly with diameters under
15 cm) increases with stand age or stand volume. In managed forests at all stages of stand development there is a
deficiency of thick deadwood. Both standing and lying deadwood at different decay stages is available contin­
uously, irrespective of the values of individual independent variables, but considerable differences exist. While
most lying deadwood exhibits higher levels of decomposition, in standing deadwood the proportions of different
decay stages are strongly associated with tree diameter at breast height. The developed models make it possible
to predict the volume of individual deadwood types for a broad range of independent variables. The current work
presents several examples, with the results showing extremely complex relationships between deadwood di­
versity and site and stand features at every stage of forest development, with continuous changes in the volume
and proportions of different deadwood types. In general, at the landscape level Polish forests contain both
standing and lying deadwood at all decay stages in more or less equal proportions. However, in forest man­
agement one should pay special attention to the dimensions of retained deadwood. The absence of thick
deadwood is particularly conspicuous in lowland managed forests.

1. Introduction saproxylic species belonging to different taxonomic groups (Stokland

A substantial proportion of forest-dwelling organisms depend on
wounded or decaying woody material derived from living, weakened, or
dead trees (Bauhus et al., 2018). Therefore, the quantity and quality of
deadwood are commonly used in nature conservation as indicators of
forest biodiversity (Venier et al., 2015; Vítková et al., 2018). The issue of
deadwood diversity is of crucial importance to saproxylic organisms
which rely on the presence of deadwood characterized by some specific
features (Andringa et al., 2019; Ramírez-Hernández et al., 2019; Nikolov
et al., 2022; Scaccini, 2022). Position, thickness, and decay stage are
very important parameters determining deadwood quality (Rimle et al.,
2017; Procházka and Schlaghamerský, 2019). The literature provides
numerous examples of relationships between these parameters and
et al., 2012). Consequently, reduced deadwood diversity often decreases
the biodiversity of saproxylic species in a given area (Pasanen et al.,
2014; Roth et al., 2019; Rieker et al., 2022) due to the lower availability
and variety of breeding and feeding niches. This prevents the occurrence
of the more specialized organisms, many of which are rare or endan­
gered (Seibold et al., 2015; Paillet et al., 2017; Ettwein et al., 2020).
Deadwood is a dynamic resource. Dead and decomposing trees arise
as a result of tree mortality, harvesting, as well as natural disturbances
(Bauhus et al., 2018). The process of decomposition alters the physical
and chemical properties of wood, which gradually turns into microsites
with different attributes (Purahong et al., 2018), leading to gradual
changes in the species composition of deadwood-dependent organisms.
A good example of this process is the succession of saproxylic insects

* Corresponding author.
E-mail address: leszek.bujoczek@urk.edu.pl (L. Bujoczek).

https://doi.org/10.1016/j.foreco.2023.121609
Received 28 April 2023; Received in revised form 18 November 2023; Accepted 20 November 2023
0378-1127/© 2023 Elsevier B.V. All rights reserved.
L. Bujoczek et al.
(Lee et al., 2014; Seibold et al., 2023). Another example is lying dead­
wood in a subalpine forest, where colonization starts with lichens, fol­
lowed by bryophytes and finally herbs and tree saplings (Zielonka and
Piątek, 2004). Standing deadwood provides a different kind of micro­
habitats, which, similarly to lying deadwood, are used by organisms in
different ways, depending on the time elapsed from tree death (e.g.,
breeding cavities, lookouts for birds of prey, feeding places) (Mikusiński
et al., 2018). Initially, snags are mostly used by saproxylic beetles and
foraging woodpeckers. Subsequently, as deadwood softens, cavities may
be made by species with weaker beaks, such as Leiopicus medius or
Dryobates minor (Pasinelli, 2007; Pakkala et al., 2019). Another very
important attribute of deadwood is its size (Heilmann-Clausen and
Christensen, 2004; Juutilainen et al., 2011; Parisi et al., 2018). Silvi­
cultural procedures substantially reduce the dimensional diversity of
dead trees, and especially limit the availability of large deadwood,
which is scarce in most managed forests (Regnery et al., 2013; Hallinger
et al., 2018; Lutz et al., 2021a; Lutz et al., 2021b). Large deadwood
ensures more stable humidity and temperature conditions for the or­
ganisms inhabiting it, enabling survival and development (Rimle et al.,
2017; Thorn et al., 2020).
The total volume of deadwood in forest ecosystems depends,
amongst others, on: forest type, stand age and density, growing stock
volume, the diameter structure of the stand, species composition, site
fertility and moisture conditions, as well as forest management (Oettel
et al., 2020; Bujoczek et al., 2021; Petritan et al., 2023). However, the
simultaneous effects of the various factors on the quantity and diversity
of deadwood have not been analyzed in great detail. Indeed, it is difficult
to comprehensively describe the processes affecting both deadwood
quality and quantity. And it is deadwood diversity that is often crucial in
the protection of saproxylic organisms.
In the current work, we set out to provide a comprehensive
description of deadwood diversity in terms of quantity and quality,
together with the factors that affect it (Fig. 1). Analysis involved three
deadwood parameters: decay stage, position, and diameter, which were
subdivided into several categories each. As a result, we distinguished 30
deadwood types on the basis of different combinations of those cate­
gories. Our aim was to address the following questions: (1) Is it possible
to estimate the volume of individual types of deadwood based on the
characteristics of forest stands and sites, and, if so, then with what de­
gree of accuracy?; (2) How does deadwood diversity vary across stands
of different ages?; (3) Does site fertility affect the volume of individual
deadwood types; (4) What are the differences in deadwood diversity
between managed and unmanaged forests?; (5) Deadwood with what
qualities is insufficiently available in forest ecosystems and what can be
done to increase deadwood diversity? This work provides guidance for
forest management, indicating deficiencies in deadwood diversity and
linking them to specific stand parameters. It also contributes to the
discussion about threshold values for deadwood volume, pointing to
differences not only in this aspect, but also in deadwood diversity. Thus,
the study may also bring economic benefits by helping to optimize
guidelines for retaining deadwood in forest ecosystems based not only
Fig. 1. Diagram of data processing rules (the 30 distinguished deadwood types are characterized in chapter 2.2).
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Forest Ecology and Management 553 (2024) 121609
on quantitative, but also qualitative, parameters.
2. Methodology
2.1. Study area and measurement data
The study used data from 29,098 sample plots distributed on a reg­
ular 4 × 4 km grid across the entire area of Poland. Individual sample
plots had an area of 0.02–0.05 ha each, depending on the age of the
dominant tree species (<60 years old – 0.02 ha, ≥61 years old – 0.04 ha,
stands subjected to selection or regeneration cuttings – 0.05 ha) (NFI,
2014). Measurements were done as part of the National Forest Inventory
(NFI) in the years 2010–2014. Living trees had their height and diameter
at breast height (DBH) taken. Only trees with DBH ≥ 7 cm were
included. Sample plots were assigned characteristics according to the
NFI methodology: terrain (lowland vs. upland or mountainous – low­
lands up to an elevation of 200–250 m a.s.l.; uplands – 200–300 m a.s.l.
including some areas above 400 m a.s.l. and elevated with respect to the
surrounding land by several dozen meters, and mountain areas – more
than 300 m a.s.l.); management type (managed forest vs. national park
or reserve); stand age based on the age of the dominant tree species
(dominant in terms of volume or area in young stands; if two or more
species had similar shares in a stand, the dominant species was defined
as that which was more desirable on a given site); site fertility (dystro­
phic/oligotrophic vs. mesotrophic/eutrophic); site moisture (dry/mesic
vs. moist/boggy). A detailed description of principles how to classify
sites according to these characteristics can be found in the NFI instruc­
tion manual (ME, 2010; NFI, 2014; Talarczyk, 2014). The studied
deadwood parameters included position (lying or standing), decay
stage, and dimensions. The length or height of deadwood was measured,
depending on its position. Entire standing and lying dead trees (with
DBH ≥ 7 cm) had their DBH taken, while diameter at mid-length (or
mid-height) was determined for tree fragments, such as logs, branches,
and snags. The study included lying trunks/logs and branches or their
fragments with a diameter at the thicker end of ≥ 10 cm and snags
(standing snapped trees) with DBH ≥ 7 cm. Stumps shorter than 0.3 m
were excluded from analysis. The decomposition of each deadwood
fragment was determined on a three-point scale: decay stage I –
non-decayed (unaltered wood structure, natural bright color, not over­
grown by fungi or bryophytes); II – partially decayed (discolored dark
wood, presence of fungi and bryophytes, rot); and III – strongly decayed
(strongly overgrown by fungi, lichens, and mosses, sapwood sometimes
completely decomposed, heartwood partially preserved) (ME, 2010;
NFI, 2014; Talarczyk, 2014).
2.2. Combining data into layers and types of deadwood
Due to the small area of individual sample plots, they were combined
into layers consisting of sample plots with the same features: manage­
ment type, terrain, site fertility, site moisture, and approximate stand
age. As a result, sample plots within a layer were characterized by a
L. Bujoczek et al.
similar species composition and silvicultural treatments. A total of 707
layers were generated. As sample plots for deadwood research should
have a minimum of 0.5–1.0 ha (Marchetti, 2005), in the present study
each layer consisted of sample plots with a combined area of approx.
1.2 ha. Stand volume (m3 ha− 1), the density of living trees (trees ha− 1),
and the mean stand age (years) were determined for each layer.
Furthermore, for each layer we calculated the volume per unit of
area (m3 ha− 1) of the 30 studied types of deadwood, characterized by
the following parameters: position (lying, L, or standing, S), decay stage
(I, II, III), and size (five classes for lying and another five for standing
deadwood). The size classes were based on DBH for standing deadwood
and on diameter for lying deadwood: 7–14.9, 15–29.9, 30–44.9,
45–59.9, and ≥ 60.0 cm. If the end diameters of a fragment of lying
deadwood were not within the same size class, such fragments (logs)
were divided into segments on the basis of a taper rate of 1 cm per 1 m,
and their length was measured. Each of the 30 types of deadwood was
characterized by a different combination of the three characteristics: L/
I/7–14 (lying at decay stage I and a diameter of 7–14.9 cm), L/I/15–29,
L/I/30–44, L/I/45–59, L/II/≥ 60, L/II/7–14, L/II/15–29,..., S/I/7–14
(standing at decay stage I and DBH of 7–14.9 cm), S/I/15–29, S/I/
30–44,..., S/III/≥ 60.
2.3. Data analysis
In the first step, the 707 layers were characterized by stand age, tree
density, stand volume, and deadwood volume (Fig. 1). In the next step,
we developed models describing deadwood diversity (the volume of
each of the 30 studied deadwood types). For that purpose, it was
necessary to use statistical tools capable of accommodating the uneven
distribution of deadwood in the field, as it is absent from the majority of
sample plots and lacks substantial diversity, especially in managed
forests. Consequently, there are many zero values in the data. The
developed model analyzed a total of 21,210 cases – this number is a
product of 707 layers multiplied by 30 deadwood types. Zeroes occurred
in as many as 12,939 cases. The independent variables used for pre­
dicting the volume of individual types of deadwood were layer charac­
teristics. The dichotomous variables were management type (managed
forest vs. national park or reserve), terrain (lowland vs. upland/moun­
tainous), while the continuous variables included stand age, stand vol­
ume, and the density of living trees (Appendix A). The site conditions
used in the generation of layers (see Section 2.2) are reflected in the
model indirectly, by the volume and density of living trees. These var­
iables are more readily available in practice, which makes the developed
model widely applicable.
Several statistical tools were tested in terms of evaluating the volume
of the 30 individual types of deadwood; these included a mixed model
with random effects and multivariate regression. Due to the specific
characteristics of the data and model quality, the hurdle model proposed
by Cragg (1971) was found to be the most suitable. In the process of
analysis, two models had to be developed for each of the 30 deadwood
types. The first one (a zero model) yielded the probability of zero counts,
while the second one (a count model) predicted the exact value of the
dependent variable for the non-zero cases. The parameters of the zero
model were estimated for the entire dataset, and the parameters of the
other one – only for the pool of those observations for which the
dependent variable was not equal to zero. The quality of the obtained
predictions was evaluated by calculating the mean absolute error
(MAE), accuracy, and area under the curve – receiver operating char­
acteristic (AUC ROC).
The developed models make it possible to process data with a wide
range of values of independent variables (Appendix A). This is in
particular true of different stand ages, volumes and densities. The
developed hurdle models can be used to calculate the volume of indi­
vidual 30 deadwood types within the broad range of values assumed by
the independent variables. To visualize changes in the volume of
different deadwood types with stand age, the diagrams present results
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Forest Ecology and Management 553 (2024) 121609
for selected ages: 0 (clear cutting area), 20, 60, 100, 130, and 140 years.
Calculations were made for the values of stand volume and stand di­
versity determined for one of the layers at a given stand age. It was the
layer for which stand volume assumed the middle value after data
ranking. Spearman’s rank correlation was used to statistically check
whether the volume of deadwood types increases or decreases with the
age of the stand. Calculations were made using hurdle models according
to stand volume, tree density and age data from all 707 layers (Fig. 2,
Appendix A, Table A1). The volume of 30 deadwood types in each layer
was calculated. Then, correlations were made between the age of the
layers and the volume of each deadwood type. The models were also
used to calculate the volume of the various deadwood types for 100-
year-old stands exhibiting different values of stand volume and tree
density. These values were both low and high, but remained within the
range observed in stands of this age (Appendix A, Table A1). Differences
were attributable to site fertility levels. All analyses were conducted for
four basic types of forests. A separate analysis was performed for low­
land managed forests and upland/mountainous managed forests. Simi­
larly, forests in national parks and reserves were divided into lowland
and upland/mountainous. Sample calculations of the volume of a
selected deadwood type are described in Appendix B.
3. Results
3.1. Layer and deadwood characteristics
Due to the high variability of the analyzed stands (707 layers), the
obtained tree density and stand volume values represent a range of
values characteristic of a given climate zone. Stands located in lowland
areas exhibit higher tree density, but lower stand volume, as compared
to those located in upland or mountainous areas. In the lowlands,
deadwood volume in the various layers of managed forests ranges from
0 to 39 m3 ha− 1, with a mean of 4 m3 ha− 1; the corresponding values for
the uplands and mountains are 1–42 m3 ha− 1 with a mean of 12 m3
ha− 1. Deadwood volume is higher in national parks and reserves and
amounts in lowlands to 5–70 m3 ha− 1 with a mean of 26 m3 ha− 1 and in
upland or mountainous areas to 21–77 m3 ha− 1 with a mean of 45 m3
ha− 1 (Fig. 2). A separate analysis of deadwood position, decay stage, and
size revealed that deadwood is highly diverse (Fig. 3) in each of the four
defined forest types. The proportion of standing deadwood is, however,
similar in all of them. Deadwood in decay stage II is the most abundant.
3.2. Changes in the volume of individual deadwood types
Both the zero model and the count model indicate the statistical
significance of independent variables, whose effects vary considerably
for different combinations of deadwood position, size, and decay stage
(a total of 30 deadwood types). The parameters of these models are
presented separately for standing deadwood (Table 1) and lying dead­
wood (Table 2). Appendix B additionally gives confidence intervals and
p-values. Depending on deadwood type, one can see certain differences
in the significance of independent variables both for standing and lying
deadwood. In most models, statistical significance obtained for both the
variable upland/mountain vs. lowland forests and the variable reserves
vs. managed forests. In the case of the zero model, parameters are
usually negative, while those for the count model are typically positive.
The other variables, i.e., stand age, volume, and tree density, are
mutually interconnected features of stands. Their statistical significance
varies between models, but each of them is significant in 22–25 models.
The quality of the developed models can be considered good
(Table 4). The obtained accuracy values range from 63.2% to 98.9%, but
in most cases they exceed 80–90%. Also satisfactory are the values of the
area under the curve (AUC), which generally amount to approx. 0.75
and more. In addition, mean absolute errors (MAE), or the mean dif­
ference between predictions and the actual values of the dependent
variable, are not high. They are expressed in the same units as the
L. Bujoczek et al.
Fig. 2. Relationships between stand age and tree density (orange points), stand volume (blue points) and deadwood volume (gray points) in the analyzed forest
types. The points represent values for each of the analyzed 707 layers.
dependent variable and amount to 0–0.35 m3 ha− 1.
The models show that the four studied forest types are markedly
different from each other in terms of deadwood diversity (reflected by
the volumes of the 30 deadwood types). Positive or negative changes in
the volumes of the various deadwood types occur with stand age (Fig. 4).
This is true of each of the four forest categories. In most cases, the vol­
ume of individual deadwood types increases with stand age, except for
thin deadwood types, whose volume tends to decrease. Spearman’s rank
correlations confirm this relationship (Table 3). Tables 1 and 2 indicate
that age was not always statistically significant in the models, but other
variables were largely related to it, i.e., the density and volume of living
trees (Fig. 2). Other factors associated with age in managed forests
include silvicultural treatments. In turn, in reserves and national parks,
depending on the stage of stand development, the dominant natural
processes are competition (in younger stands) and tree mortality (in
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Forest Ecology and Management 553 (2024) 121609
older stands). The volume of individual deadwood types does not exceed
4 m3 ha− 1, with the highest values recorded in upland/mountain na­
tional parks and reserves. At the other end of the spectrum, the corre­
sponding value for lowland managed forests typically ranges from 0 to
0.4 m3 ha− 1, with a maximum of 0.7 m3 ha− 1. Those forests are also
characterized by very low quantities of deadwood with dimensions
exceeding 45 cm, even in stands aged 100 and 130 years. This is
particularly true of lying deadwood, where the volume of deadwood
types of such dimensions tends to be around 0.1 m3ha-1.
Fig. 5 illustrates the effects of stand volume and tree density in 100-
year-old stands. These variables are often statistically significant in the
developed models. Differences in stand volume are mostly attributable
to site fertility. The values obtained after entering independent variables
in hurdle models show that the volume of almost all deadwood types
increases in stands that are of the same age but have a higher stand
L. Bujoczek et al.
Fig. 3. Mean deadwood volume depending on deadwood position, decay stage, and size.
volume.
4. Discussion
This study presents a complex picture of changes in the quantity of
different deadwood types in terms of forest development stages and
deadwood features determined by site conditions and management
treatments. The findings also make it possible to understand deficiencies
in deadwood diversity in terms of individual deadwood types rather
than its total volume. It should be noted that multiple deadwood types,
but in different quantities, were found at almost every stage of forest
development, which is due to the fact that at any stand age deadwood
consists partly of fresh wood (from trees that died recently) and old
deadwood that has been there for a long time. This is particularly pro­
nounced in young stands featuring some large-diameter deadwood left
over from past cuttings (Lombardi et al., 2008). The residence time of
different deadwood fragments in the stand mostly depends on the rate of
decomposition, which is affected by numerous factors (Přívětivý et al.,
2016; Gómez-Brandón et al., 2020; Jomura et al., 2022). All of these
factors, together with natural disturbances and management treatments,
will determine the proportions of the various deadwood types in a given
area.
The method of data analysis was based on layers constructed from
multiple NFI sample plots, due to which our results are valid at the
landscape level. This is significant as individual stands with small areas
may in reality greatly differ from one another in terms of deadwood
quantity and quality. Thus, evaluation at the stand level may indicate
low deadwood diversity. For instance, an examination of an even-aged
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Forest Ecology and Management 553 (2024) 121609
single-story monospecific stand locally affected by a violent abiotic
factor (e.g., gale) or a biotic factor (e.g., insect outbreak) may reveal
relatively homogeneous deadwood. In such a case, deadwood position,
decay stage, and size may be dominated by one feature or value. By
extending the inventory area to the forest or landscape level, one will
arrive at results consistent with those obtained from the models pre­
sented in this work. Furthermore, these larger scales seem to be more
appropriate for the analysis of deadwood diversity. They are better
suited to describing the pattern of silvicultural treatments in managed
forests and the resulting deadwood diversity, rather than random dis­
turbances affecting individual stands. In such stands, deadwood result­
ing from random events is usually removed as part of sanitary or salvage
cuttings within a short period of time. Many saproxylic organisms have
the ability to move and colonize new areas (Jusino et al., 2016;
Johansson et al., 2021), but relatively little is known about this phe­
nomenon, at least for some taxonomic groups (Komonen and Müller,
2018; Nordén et al., 2018; Martin at al, 2021). The literature indicates
both positive and negative aspects of irregular deadwood distribution
within a landscape (Augustynczik, 2021; Haeler at al, 2021; Mazziotta
et al., 2023), both in terms of ecological and economic considerations.
With a view to the protection of saproxylic organisms, it is important for
areas with favorable habitat conditions not to be separated islands,
which would prevent species expansion or genetic exchange.
This work shows that many distinct deadwood types occur in low
volumes per unit of area. Thus, the question arises as to whether silvi­
cultural guidelines should primarily focus on scarce deadwood types
with threshold values specified for them. Another question is whether
some deadwood types are continuously present in stands irrespective of
L. Bujoczek et al. Forest Ecology and Management 553 (2024) 121609

Table 1
Zero model and count model parameters for standing deadwood.

1
Deadwood type – S – standing; I, II, III – decay stage; 7–14, 15–29, 30–44, 45–59, ≥ 60 – diameter [cm].
The upper rows (blue figures) represent the parameters of the zero model, while the bottom ones (black figures) pertain to the count model. * – p < 0.05; * * – p < 0.01;
* ** – p < 0.001.

management practices. Both questions may be answered in the affir­
mative. Research shows that a more detailed approach is justified both
from the environmental and economic standpoints (Vítková et al.,
2018). The enrichment of forest ecosystems should concentrate on
several selected characteristics of deadwood, such as various decay
degrees, deadwood tree species and deadwood size (Blaser, Öder et al.,
2013, 2021; Yang et al., 2021). We also believe that deadwood char­
acteristics can be divided into those that may be to some extent
controlled, and those which are naturally diverse, without any in­
terventions, once a sufficient quantity of deadwood is accumulated (on a

6
L. Bujoczek et al. Forest Ecology and Management 553 (2024) 121609

Table 2
Zero model and count model parameters for lying deadwood.

1
Deadwood type – L – lying; I, II, III – decay stages; 7–14, 15–29, 30–44, 45–59, ≥ 60 – diameter [cm]. The upper rows (blue figures) represent the parameters of the
zero model, while the bottom ones (black figures) pertain to the count model. * – p < 0.05; * * – p < 0.01; * ** – p < 0.001.

forest or landscape level). This is consistent with other studies, which
have found a positive relationship between deadwood volume and di­
versity (Haeler at al, 2021; Bujoczek and Bujoczek, 2022).
The present work analyzed three characteristics of deadwood. The
proportions between standing and lying deadwood were found to be
quite even. They are influenced by numerous factors, with the main ones
being the cause of tree death, wind, terrain, and tree species (Csilléry
et al., 2017; Hotta et al., 2021; Merganič et al., 2022). Some in­
terventions to shape these proportions may be undertaken locally, but
only in exceptional cases. Also decay stage is a feature that does not

7
L. Bujoczek et al.
Fig. 4. Volumes (values calculated based on hurdle models) of 30 individual deadwood types by stand age (the stand volume and tree density values used for
calculations are contained in Appendix A).
require human interference. Research has shown that the retention of all
decay classes of deadwood on a landscape scale is necessary to conserve
saproxylic fauna (Lee et al., 2014). The present study found substantial
diversity in terms of deadwood classes on the analyzed spatial scale.
Interventions may only be needed with regard to some local populations
of endangered saproxylic species. Furthermore, it should be noted that
proportions between decay stages differ between standing and lying
deadwood (Oettel et. al, 2020), which is a natural phenomenon attrib­
utable to the loss of stability of standing dead trees due to gradual
decomposition. At the time of falling, standing deadwood is often at an
advanced stage of decay, and so the proportion of fresh lying deadwood
is lower. On the other hand, some rot fungi and fungal fruiting bodies
can be also found on living trees, and especially those with a large DBH.
As can be seen, decay stage is affected by multiple factors, ensuring
diversity (Jomura et al., 2022). While tree species differ considerably in
their rates of decomposition, there are many variables that may slow
down or accelerate this process (Hararuk et al., 2020). An important role
is played by site and climatic conditions, the cause of tree death, frag­
mentation upon downing, the drying of wood, and the time over which
the tree remains standing after death (Vrška et al., 2015; Chang et al.,
2020; Kipping et al., 2022).
Species composition and size are those aspects that exert a
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Forest Ecology and Management 553 (2024) 121609
substantial effect on deadwood diversity and lend themselves to
enrichment. The former feature was not addressed in the present study,
but it is nevertheless crucial due to the species specificity of some sap­
roxylic organisms (Parisi et al., 2018). Another issue is the size structure
of deadwood. It is an important criterion in evaluating the conservation
state of Natura 2000 sites, for which a minimum number of dead trees is
often specified (Alberdi et al., 2019). It should be noted that the state of
many sites is classified as unfavorable-inadequate or unfavorable-bad
(Bujoczek et al., 2020), amongst others due to an insufficient avail­
ability of suitable deadwood. Fine woody debris does play a certain role
in the ecosystem, but only for some saproxylic organisms, and especially
fungi (Heilmann-Clausen and Christensen, 2004; Piętka et al., 2019). In
general, such deadwood is present both in managed and unmanaged
forests.
This work focused on the analysis of deadwood volume. Even if the
volume of individual fragments of thin deadwood is small, its overall
amount is comparatively high. Our own field observations as well as the
size structure of standing deadwood indicate that this finding results
from the large number of branches and thin dead trees. In addition, they
occupy a large area on the forest floor relative to their volume. The
absence of the oldest and thickest trees is most conspicuous in managed
lowland forests, mostly due to the adopted rotation age regime.
L. Bujoczek et al. Forest Ecology and Management 553 (2024) 121609

Table 3 Table 4
Spearman’s correlation between the age and volume of deadwood species Evaluation of model quality.
(calculations of deadwood volume based on hurdle models for the values of Deadwood type1 Accuracy Area under Mean absolute error
stand volume, tree density and age in 707 layers). the curve
Deadwood Lowland Upland/ Lowland Upland/ L/I/7–14 82.32% 0.683 0.0648
type1 managed mountainous national mountainous L/I/15–29 63.22% 0.683 0.1905
forests managed parks and national parks L/I/30–44 80.48% 0.708 0.0996
forests reserves and reserves L/I/45–59 94.63% 0.818 0.0115
S/I/7–14 -0,67* -0,78* -0,97* -0,94* L/I/> 60 98.87% 0.885 0.0014
S/I/15–29 0,24* 0,42* -0,27 0,15 L/II/7–14 91.09% 0.815 0.1046
S/I/30–44 0,87* 0,89* 0,71* 0,79* L/II/15–29 75.53% 0.734 0.2265
S/I/45–59 0,94* 0,95* 0,92* 0,95* L/II/30–44 76.94% 0.769 0.0882
S/I/> 60 0,91* 0,86* 0,95* 0,60 L/II/45–59 90.95% 0.787 0.0348
S/II/7–14 -0,65* -0,77* -0,96* -0,82* L/II/> 60 97.45% 0.768 0.0338
S/II/15–29 0,29* 0,15 -0,36 -0,06 L/III/7–14 91.37% 0.774 0.104
S/II/30–44 0,78* 0,89* 0,58* 0,85* L/III/15–29 83.88% 0.782 0.2653
S/II/45–59 0,76* 0,78* 0,95* 0,43 L/III/30–44 74.68% 0.771 0.1355
S/II/> 60 0,93* 0,97* 0,96* 0,97* L/III/45–59 86.99% 0.774 0.0467
S/III/7–14 -0,13* -0,42* -0,71* -0,87* L/III/≥ 60 94.63% 0.846 0.0324
S/III/15–29 -0,05 -0,10 -0,56* -0,87* S/I/7–14 91.65% 0.828 0.0849
S/III/30–44 0,93* 0,95* 0,86* 0,95* S/I/15–29 82.04% 0.773 0.3511
S/III/45–59 1,00* 1,00* 0,99* 0,97* S/I/30–44 73.69% 0.75 0.2951
S/III/≥ 60 0,64* 0,68* 0,96* 0,76* S/I/45–59 93.49% 0.763 0.0986
L/I/7–14 -0,71* -0,90* -1,00 -1,00 S/I/> 60 97.88% 0.853 0.028
L/I/15–29 0,63* 0,96* 0,23 0,83* S/II/7–14 94.77% 0.849 0.1296
L/I/30–44 0,93* 0,96* 0,90* 0,95* S/II/15–29 86.85% 0.79 0.3473
L/I/45–59 0,99* 0,95* 0,87* 0,86* S/II/30–44 70.72% 0.77 0.3408
L/I/> 60 0,25* 0,89* 0,87* 0,96* S/II/45–59 91.94% 0.848 0.0942
L/II/7–14 -0,48* -0,52* -0,88* -0,61* S/II/> 60 95.90% 0.846 0.0467
L/II/15–29 0,61* 0,77* 0,36 0,69* S/III/7–14 70.72% 0.736 0.0419
L/II/30–44 0,88* 0,90* 0,83* 0,85* S/III/15–29 67.04% 0.731 0.1043
L/II/45–59 0,53* 0,66* 0,51* 0,65* S/III/30–44 80.34% 0.771 0.0732
L/II/> 60 0,63* 0,78* 0,47 0,63* S/III/45–59 93.35% 0.84 0.0116
L/III/7–14 -0,18* -0,54* -0,86* -0,94* S/III/≥ 60 96.04% 0.845 0.0156
L/III/15–29 0,77* 0,86* 0,57* 0,79* 1
Deadwood type – L – lying, S – standing; I, II, III – decay stages; 7–14, 15–29,
L/III/30–44 0,98* 0,99* 0,96* 0,99*
30–44, 45–59, ≥ 60 – diameter [cm].
L/III/45–59 0,87* 0,91* 0,89* 0,85*
L/III/≥ 60 0,74* 0,80* 0,69* 0,80*
1 possibilities of increasing deadwood volume for economic reasons.
Deadwood type – L – lying, S – standing; I, II, III – decay stages; 7–14, 15–29,
30–44, 45–59, ≥ 60 – diameter [cm]. Addressing the question posed in the title, we found that stands differ
*
Results are statistically significant at p < 0.05. considerably not only in terms of deadwood quantity, but also diversity.
The quantities of the various deadwood types differed between stands of
Deadwood with a diameter of more than 30 cm is scarce, and that with a different ages, characterized by different stand volumes and tree den­
diameter greater than 45 cm is almost entirely absent. More deadwood sities. In general, the higher the stand age and the more fertile the forest
(also large) is found in managed upland/mountain forests because of site (factors associated with stand volume) the higher the volumes of
higher rotation ages as compared to pine, which is predominant in individual deadwood types, which are also typically greater in unman­
lowlands, as well as owing to more fertile sites and greater terrain- aged forests. Among the three deadwood characteristics studied in this
related difficulty of management. As a result, some dead trees remain work, position and decay stage exhibit high diversity on the landscape
unnoticed or uneconomical to harvest, especially that wood is subject to level and do not necessitate human interference. On the other hand, the
depreciation after some time (Bujoczek et al., 2021). Thus, in easily size diversity of deadwood is insufficient, especially in managed forests,
accessible terrain with low availability of large deadwood it is impera­ where mostly thin deadwood is found. The retention of several large
trees until natural death per hectare will enrich both deadwood diversity
tive to designate areas or trees excluded from harvesting. Some efforts to
increase the quantity and diversity of deadwood have been already and increase total deadwood volume in managed forests, which is
generally low. In addition, the present work shows differences in the
implemented and should improve the size structure of deadwood in the
long term. Beginning several years ago, 5% of the area of harvested volume of individual deadwood types at different stages of forest
development. This is natural and observed also in national parks and
stands is to be retained. Trees in the designated areas will have an op­
portunity to grow to large dimensions and will be left in place until they reserves. Therefore, any threshold values for different deadwood types
should take into account stand and site characteristics. Forests in na­
eventually die and decompose (IOL, 2012; Referowska-Chodak, 2014).
The quality and consistency of this practice will determine changes in tional parks and reserves may provide a reference standard for pro­
portions between different deadwood types.
deadwood quantity and quality in the coming decades.

CRediT authorship contribution statement

5. Conclusions
Bujoczek Leszek: Conceptualization, Formal analysis, Methodol­
The main objective of the present study on deadwood availability
ogy, Visualization, Writing – original draft, Writing – review & editing.
was to determine those types of deadwood that are deficient in the
Bujoczek Małgorzata: Visualization, Writing – review & editing. Zięba
ecosystem in the context of the conservation of saproxylic organisms,
Stanisław: Visualization, Writing – review & editing.
and consequently to make it possible to remedy those deficiencies. It has
been increasingly suggested that the criterion of total deadwood quan­
tity is insufficient. Even though literature reports indicate that there is a Declaration of Competing Interest
positive relationship between deadwood quantity and diversity,
managed forests require more precise interventions due to the limited The authors declare that they have no known competing financial

9
L. Bujoczek et al. Forest Ecology and Management 553 (2024) 121609

Fig. 5. Comparison of the volumes of individual deadwood types (values calculated based on hurdle models) in 100-year-old stands with different stand volumes and
tree densities (gray – lower stand volume at higher tree density; orange – higher stand volume at lower tree density. The exact stand volume and tree density values
used for calculations are given in Appendix A).

interests or personal relationships that could have appeared to influence Acknowledgments

the work reported in this paper.
This study was financed by the Ministry of Science and Higher Ed­
Data Availability ucation of the Republic of Poland. The authors thank the State Forests
National Forest Holding for making the data available, as well as the
This data is part of the National Forest Inventory and has been made anonymous Reviewers for their comments.
available to the authors for this publication. They are owned by the State
Forests National Forest Holding.

Appendix A
Table A1
Characteristics of the 707 analyzed layers (A mean stand age was calculated for each layer; in the table the layers were assigned to rows according to the age range
specified in the “stand age” column.).

Stand age [years] Stand volume ean (min–max) [m3 ha–1] Tree density Mean (min–max) [trees ha–1] Deadwood volume Mean (min–max) [m3 ha–1]
Managed lowland forests

0 0 (0–0) 0 (0–1) 3 (0–12)

1–10 16 (5–57) 88 (5–349) 1 (0–6)
11–20 52 (11–100) 1114 (357–1932) 1 (0–4)
21–30 132 (88–164) 1472 (1042–1928) 2 (1–6)
31–40 199 (153–257) 1190 (796–1644) 5 (1–16)
41–50 267 (216–340) 995 (654–1427) 4 (1–12)
(continued on next page)

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L. Bujoczek et al. Forest Ecology and Management 553 (2024) 121609

Table A1 (continued )
Stand age [years] Stand volume ean (min–max) [m3 ha–1] Tree density Mean (min–max) [trees ha–1] Deadwood volume Mean (min–max) [m3 ha–1]
Managed lowland forests

51–60 298 (258–341) 831 (594–1395) 5 (1–28)

61–70 310 (244–395) 663 (520–927) 4 (1–16)
71–80 342 (255–446) 641 (504–833) 5 (1–39)
81–90 359 (259–476) 588 (505–686) 4 (0–17)
91–100 368 (287–508) 531 (444–608) 5 (1–19)
101–110 389 (311–482) 492 (352–576) 6 (1–34)
111–120 400 (330–495) 460 (382–577) 6 (1–17)
121–130 429 (360–538) 447 (382–488) 5 (1–10)
> 130 448 (351–567) 401 (340–514) 10 (3–19)
Managed upland/mountain forests
1–10 34 (8–67) 152 (13–356) 7 (2–13)
11–20 72 (64–85) 648 (249–891) 7 (4–11)
21–30 121 (108–134) 1008 (903–1069) 6 (4–8)
31–40 192 (168–218) 957 (932–1005) 9 (5–11)
41–50 241 (202–266) 895 (769–1034) 12 (6–19)
51–60 301 (248–357) 753 (624–929) 12 (6–22)
61–70 335 (265–440) 628 (492–798) 12 (1–27)
71–80 365 (300–465) 568 (478–642) 11 (5–26)
81–90 405 (302–454) 542 (474–636) 11 (4–25)
91–100 439 (316–490) 478 (360–638) 13 (6–22)
101–110 448 (376–521) 440 (331–601) 14 (6–22)
111–120 459 (416–528) 407 (346–503) 11 (8–13)
121–130 472 (442–508) 342 (253–431) 20 (7–42)
> 130 494 (451–536) 317 (301–332) 24 (18–29)
Lowland forests in national parks and reserves
31–40 221 (180–261) 896 (813–980) 17 (17–17)
41–50 204 (197–210) 1094 (893–1296) 15 (5–25)
61–70 335 (296–387) 728 (670–771) 25 (8–34)
81–90 390 (363–426) 580 (549–615) 37 (5–70)
101–110 327 (327–327) 696 (696–696) 26 (26–26)
111–120 412 (382–468) 526 (511–547) 33 (8–69)
> 130 446 (371–521) 487 (467–506) 19 (14–23)
Upland/mountain forests in national parks and reserves
21–30 121 (92–149) 691 (539–844) 49 (21–77)
51–60 352 (352–352) 844 (844–844) 50 (50–50)
61–70 409 (409–409) 683 (683–683) 31 (31–31)
81–90 429 (429–429) 737 (737–737) 26 (26–26)
91–100 527 (527–527) 558 (558–558) 34 (34–34)
101–110 571 (571–571) 496 (496–496) 41 (41–41)
111–120 441 (350–532) 517 (442–592) 54 (35–73)
121–130 578 (578–578) 380 (380–380) 38 (38–38)
> 130 648 (648–648) 411 (411–411) 67 (67–67)

Values given in Fig. 4 (Manuscript).

Volumes in Fig. 4 were calculated based on hurdle models for the parameters presented in Tables 1 and 2 (Manuscript). The stand volume and tree
density values given in brackets were obtained for one of the layers of a given age (Appendix A, Table A1). For those layers, stand volume assumed the
middle value after data ranking. Calculations were done for the following values: managed lowland forests – 0 years, (stand volume – 0 m3 ha− 1, tree
density – 0 trees ha− 1), 20 years (94, 1487), 60 years (303, 703), 100 years (378, 508), 130 years (434, 432); managed upland/mountain forests – 20
years (90, 911), 60 years (318, 663), 100 years (425, 452), 130 years (459, 303); lowland forests in national parks and reserves – 20 years (111, 1205),
60 years (293, 810), 100 years (401, 569), 140 years (434, 483); upland/mountain forests in national parks and reserves – 20 years (115, 745), 60
years (341, 687), 100 years (500, 576), 140 years (592, 413).
Values given in Fig. 5 (Manuscript).
Volumes in Fig. 5 were calculated based on hurdle models for the parameters presented in Tables 1 and 2 (Manuscript). The stand volume and tree
density values given in brackets were obtained for one of the layers containing 100-year-old trees (Appendix A, Table A1). These were layers with the
lowest and highest stand volume values. Calculations were done for the following values: managed lowland forests – gray (stand volume 326 m3 ha− 1,
tree density 590 trees ha− 1) and orange (455, 460); managed upland/mountain forests – gray (315, 594) and orange (521, 331); lowland forests in
national parks and reserves – gray (326, 695) and orange (467, 510); upland/mountain forests in national parks and reserves – gray (350, 590) and
orange (531, 442).

Appendix B

Example of calculations for deadwood type – S/I/15–29.

Adopted values of independent variables:

1. Terrain: upland/mountainous
2. Type of management: national parks and reserves
3. Tree density – 844 trees ha− 1
4. Stand volume – 352 m3 ha− 1
5. Stand age – 55 years

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L. Bujoczek et al. Forest Ecology and Management 553 (2024) 121609

A. Zero model – likelihood that the dependent variable is zero (no deadwood).

Parameter values are given in parentheses (see Table B1 below):

1. National parks and reserves (− 1.479)

2. Uplands/mountains (− 0.7486)
3. Tree density (− 0.0002)
4. Stand volume (− 0.0198)
5. Stand age (0.0513)
6. Constant (1.231)

1.231 + 0.0513 ⋅ 55 – 0.0198 ⋅ 352 – 0.0002 ⋅ 844 – 0.7486 – 1.479 = − 5.3135.

The zero model is a logistic regression, where likelihood is computed from the formula:
ex
p=
1 + ex

where x is the value calculated above – a linear predictor.

e− 5.3135
p=
1 + e− 5.3135
The result is p = 0.0049; (0.49%).

B. Count model

Parameter values are given in parentheses (see Table B2 below):

1. National parks and reserves (1.0153)

2. Uplands/mountains (0.244)
3. Tree density (0.0003)
4. Stand volume (0.0014)
5. Stand age (0.006)
6. Constant (0.8667)

0.8667 – 0.006 ⋅ 55 + 0.0014 ⋅ 352 – 0.0003 ⋅ 844 + 0.244 + 1.0153 = 2.0356.

C. Volume S/I/15–29

In summary, the model predicts that deadwood will be absent at a probability of 0.49%, and if it is present, then its mean volume will be
2.0356 m3 ha− 1.
Which results in a final mean prediction of.
2.0356 ⋅ (1 – 0.0049) = 2.026 m3 ha-1.
Volume S/I/15–29 ¼ 2.026 m3 ha− 1 (in upland/mountainous national parks and reserves).

Table B1
Zero model – a model parameter is given for each independent variable with a confidence interval (in brackets) and a p value. Asterisks indicate statistically significant
effects (p < 0.05). Deadwood type – L – lying, S – standing; I, II, III – decay stages; 7–14, 15–29, 30–44, 45–59, ≥ 60 – diameter [cm].

Deadwood Constant Stand age [years] Stand volume [m3 Tree density Terrain National parks and
type ha¡1] [trees ha¡1] (upland/mountainous reserves vs. managed
vs. lowland forests) forests

S/I/7–14 2.1988 (1.1429; 0.0274 (0.0091; -0.0127 (− 0.018; -0.0037 (− 0.0051; -1.8963 (− 3.2216; -0.8375 (− 2.9948; 1.3198).
3.2547). p < 0.001 * 0.0456). p = 0.003 * − 0.0074). − 0.0024). p < 0.001 * − 0.5709). p = 0.005 * p = 0.447
p < 0.001 *
S/I/15–29 1.231 (0.4385; 0.0513 (0.0366; -0.0198 (− 0.0239; -0.0002 (− 0.0008; -0.7486 (− 1.4406; -1.479 (− 3.1546; 0.1966).
2.0236). p = 0.002 * 0.0661). p < 0.001 * − 0.0157). 0.0004). p = 0.528 − 0.0567). p = 0.034 * p = 0.084
p < 0.001 *
S/I/30–44 1.7948 (0.8203; 0.0065 (− 0.0063; -0.0071 (− 0.0107; 0.0017 (0.0008; -0.7525 (− 1.2113; -0.8372 (− 1.725; 0.0506).
2.7693). p < 0.001 * 0.0193). p = 0.317 − 0.0036). 0.0027). p < 0.001 * − 0.2937). p = 0.001 * p = 0.065
p < 0.001 *
S/I/45–59 2.1303 (0.8921; 0.0032 (− 0.0177; -0.0031 (− 0.0089; 0.0026 (0.001; -0.8257 (− 1.5429; -1.1305 (− 2.1892;
3.3685). p = 0.001 * 0.0241). p = 0.763 0.0026). p = 0.287 0.0043). p = 0.002 * − 0.1085). p = 0.024 * − 0.0718). p = 0.036 *
S/I/> 60 6.4419 (− 0.0059; -0.0047 (− 0.0412; -0.0105 (− 0.0219; 0.0031 (− 0.0036; 0.018 (− 1.4414; 1.4775). -0.2694 (− 2.2427; 1.704).
12.8898). p = 0.05 0.0319). p = 0.803 0.0009). p = 0.072 0.0098). p = 0.369 p = 0.981 p = 0.789
S/II/7–14 1.3626 (0.4699; 0.0207 (− 0.004; -0.0127 (− 0.0195; -0.0027 (− 0.0039; -2.4312 (− 4.5411; -15.0359 (− 2178.0183;
2.2553). p = 0.003 * 0.0453). p = 0.1 − 0.0058). − 0.0016). p < 0.001 * − 0.3213). p = 0.024 * 2147.9466). p = 0.989
p < 0.001 *
(continued on next page)

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L. Bujoczek et al. Forest Ecology and Management 553 (2024) 121609

Table B1 (continued )
Deadwood Constant Stand age [years] Stand volume [m3 Tree density Terrain National parks and
type ha¡1] [trees ha¡1] (upland/mountainous reserves vs. managed
vs. lowland forests) forests

S/II/15–29 1.0192 (0.2254; 0.0355 (0.0191; -0.0171 (− 0.0216; 0 (− 0.0006; 0.0006). -2.5292 (− 3.9949; -15.4208 (− 1228.1434;
1.813). p = 0.012 * 0.052). p < 0.001 * − 0.0126). p = 0.943 − 1.0635). p = 0.001 * 1197.3018). p = 0.98
p < 0.001 *
S/II/30–44 1.0167 (0.1988; 0.0191 (0.0061; -0.0093 (− 0.0128; 0.0021 (0.0013; -1.5246 (− 2.0075; -1.7566 (− 2.7973; − 0.716).
1.8346). p = 0.015 * 0.0321). p = 0.004 * − 0.0057). 0.0029). p < 0.001 * − 1.0416). p < 0.001 * p = 0.001 *
p < 0.001 *
S/II/45–59 3.5345 (1.6351; -0.0207 (− 0.0394; -0.0016 (− 0.007; 0.0025 (0.0004; -1.3972 (− 2.0569; -2.7558 (− 3.7646;
5.4339). p < 0.001 * − 0.0021). 0.0038). p = 0.569 0.0046). p = 0.021 * − 0.7375). p < 0.001 * − 1.7469). p < 0.001 *
p = 0.029 *
S/II/> 60 3.44 (1.2286; -0.0147 (− 0.038; -0.0028 (− 0.0099; 0.0041 (0.001; -0.5351 (− 1.544; -1.9825 (− 3.132; − 0.833).
5.6515). p = 0.002 * 0.0085). p = 0.214 0.0044). p = 0.448 0.0071). p = 0.009 * 0.4737). p = 0.299 p = 0.001 *
S/III/7–14 1.9839 (1.0973; 0.0207 (0.0079; -0.0086 (− 0.012; -0.0019 (− 0.0027; -1.4416 (− 2.0586; -2.5011 (− 4.5696;
2.8706). p < 0.001 * 0.0334). p = 0.001 * − 0.0052). − 0.0012). p < 0.001 * − 0.8246). p < 0.001 * − 0.4327). p = 0.018 *
p < 0.001 *
S/III/15–29 0.9803 (0.2176; 0.0246 (0.0119; -0.0082 (− 0.0116; 0.0004 (− 0.0002; -2.0407 (− 2.6085; -2.6886 (− 4.2189;
1.7431). p = 0.012 * 0.0373). p < 0.001 * − 0.0049). 0.001). p = 0.214 − 1.4729). p < 0.001 * − 1.1582). p = 0.001 *
p < 0.001 *
S/III/30–44 2.6981 (1.4589; 0.0069 (− 0.0073; -0.0071 (− 0.011; 0.0011 (0; 0.0022). -1.1635 (− 1.6355; -2.0279 (− 2.97; − 1.0857).
3.9373). p < 0.001 * 0.0211). p = 0.34 − 0.0031). p = 0.061 − 0.6914). p < 0.001 * p < 0.001 *
p < 0.001 *
S/III/45–59 4.0985 (2.1094; -0.0155 (− 0.0366; 0.0007 (− 0.005; 0.0007 (− 0.0012; -2.2865 (− 2.994; -2.232 (− 3.2073; − 1.2567).
6.0875). p < 0.001 * 0.0055). p = 0.149 0.0064). p = 0.814 0.0025). p = 0.48 − 1.579). p < 0.001 * p < 0.001 *
S/III/≥ 60 3.1775 (1.475; -0.0191 (− 0.0439; 0.004 (− 0.0028; 0.0018 (− 0.0002; -1.9772 (− 2.8922; -2.0812 (− 3.1675; − 0.995).
4.88). p < 0.001 * 0.0058). p = 0.133 0.0109). p = 0.247 0.0038). p = 0.083 − 1.0623). p < 0.001 * p < 0.001 *
L/I/7–14 0.692 (− 0.0447; 0.0107 (− 0.0037; -0.0063 (− 0.0101; -0.0011 (− 0.0018; -1.5905 (− 2.4565; -15.716 (− 1394.3129;
1.4287). p = 0.066 0.025). p = 0.145 − 0.0025). − 0.0005). p = 0.001 * − 0.7244). p < 0.001 * 1362.8809). p = 0.982
p = 0.001 *
L/I/15–29 1.1311 (0.3802; 0.021 (0.0092; -0.0088 (− 0.012; -0.0001 (− 0.0006; -1.2765 (− 1.8052; -0.6915 (− 1.6872; 0.3041).
1.8819). p = 0.003 * 0.0329). p = 0.001 * − 0.0057). 0.0005). p = 0.864 − 0.7478). p < 0.001 * p = 0.173
p < 0.001 *
L/I/30–44 1.7475 (0.7818; 0.0055 (− 0.0081; -0.0049 (− 0.0086; 0.0015 (0.0005; -0.8763 (− 1.3412; -0.7403 (− 1.608; 0.1274).
2.7132). p < 0.001 * 0.0191). p = 0.43 − 0.0012). 0.0024). p = 0.003 * − 0.4113). p < 0.001 * p = 0.095
p = 0.009 *
L/I/45–59 3.416 (1.384; 0.0083 (− 0.0149; -0.0084 (− 0.015; 0.0035 (0.001; 0.006). -1.0702 (− 1.8484; -1.2862 (− 2.4394;
5.448). p = 0.001 * 0.0315). p = 0.482 − 0.0018). p = 0.006 * − 0.292). p = 0.007 * − 0.1331). p = 0.029 *
p = 0.013 *
L/I/> 60 9.0095 (2.344; -0.0658 (− 0.1104; 0.01 (− 0.005; 0.025). -0.0022 (− 0.0067; -2.8994 (− 4.6698; 16.3465 (− 3478.5552;
15.675). p = 0.008 * − 0.0213). p = 0.191 0.0022). p = 0.327 − 1.129). p = 0.001 * 3511.2482). p = 0.993
p = 0.004 *
L/II/7–14 1.1099 (0.2737; 0.0279 (0.0092; -0.0125 (− 0.0177; -0.0022 (− 0.0032; -2.7539 (− 4.7897; -14.9004 (− 1298.0354;
1.9461). p = 0.009 * 0.0465). p = 0.003 * − 0.0072). − 0.0012). p < 0.001 * − 0.7182). p = 0.008 * 1268.2345). p = 0.982
p < 0.001 *
L/II/15–29 0.6988 (− 0.0365; 0.0222 (0.0089; -0.01 (− 0.0136; 0 (− 0.0006; 0.0005). -2.3707 (− 3.4018; -16.1082 (− 1285.6614;
1.4341). p = 0.063 0.0355). p = 0.001 * − 0.0065). p = 0.863 − 1.3396). p < 0.001 * 1253.445). p = 0.98
p < 0.001 *
L/II/30–44 1.3054 (0.4374; 0.0016 (− 0.0116; -0.0042 (− 0.0078; 0.0019 (0.001; -1.6326 (− 2.0962; -1.6093 (− 2.5636; − 0.655).
2.1734). p = 0.003 * 0.0147). p = 0.816 − 0.0006). 0.0028). p < 0.001 * − 1.169). p < 0.001 * p = 0.001 *
p = 0.021 *
L/II/45–59 1.6038 (0.5733; 0.015 (− 0.0052; -0.005 (− 0.0104; 0.0028 (0.0014; -1.3693 (− 1.9723; -1.8516 (− 2.7947;
2.6342). p = 0.002 * 0.0352). p = 0.146 0.0004). p = 0.068 0.0042). p < 0.001 * − 0.7663). p < 0.001 * − 0.9085). p < 0.001 *
L/II/> 60 3.5511 (1.4305; 0.0114 (− 0.0237; -0.0045 (− 0.0135; 0.0021 (− 0.0004; -1.4203 (− 2.4888; -1.0928 (− 2.5483; 0.3628).
5.6718). p = 0.001 * 0.0465). p = 0.523 0.0046). p = 0.333 0.0046). p = 0.105 − 0.3517). p = 0.009 * p = 0.141
L/III/7–14 0.774 (0.0056; 0.0315 (0.0125; -0.0151 (− 0.0203; -0.001 (− 0.0017; -1.4773 (− 2.7122; -14.9511 (− 1311.1332;
1.5424). p = 0.048 * 0.0506). p = 0.001 * − 0.0099). − 0.0003). p = 0.007 * − 0.2425). p = 0.019 * 1281.2309). p = 0.982
p < 0.001 *
L/III/15–29 0.8369 (0.0765; 0.0202 (0.0042; -0.0125 (− 0.0167; 0 (− 0.0006; 0.0006). -2.1836 (− 3.3895; -15.3775 (− 1283.8687;
1.5973). p = 0.031 * 0.0362). p = 0.013 * − 0.0082). p = 0.939 − 0.9777). p < 0.001 * 1253.1136). p = 0.981
p < 0.001 *
L/III/30–44 1.7487 (0.8428; 0.0007 (− 0.012; -0.004 (− 0.0075; 0.0007 (− 0.0001; -2.1014 (− 2.6008; -2.1846 (− 3.3298;
2.6546). p < 0.001 * 0.0135). p = 0.91 − 0.0006). 0.0015). p = 0.075 − 1.6021). p < 0.001 * − 1.0393). p < 0.001 *
p = 0.021 *
L/III/45–59 3.3221 (1.9041; -0.0036 (− 0.02; -0.0024 (− 0.0068; 0.0002 (− 0.001; -1.9002 (− 2.4089; -0.9976 (− 1.9225;
4.74). p < 0.001 * 0.0128). p = 0.666 0.0019). p = 0.276 0.0014). p = 0.765 − 1.3915). p < 0.001 * − 0.0726). p = 0.035 *
L/III/≥ 60 1.9568 (0.7409; 0.008 (− 0.0159; -0.0049 (− 0.0116; 0.0044 (0.0023; -1.4784 (− 2.2629; -1.5862 (− 2.7081;
3.1726). p = 0.002 * 0.0319). p = 0.512 0.0018). p = 0.15 0.0065). p < 0.001 * − 0.694). p < 0.001 * − 0.4644). p = 0.006 *

13
L. Bujoczek et al. Forest Ecology and Management 553 (2024) 121609

Table B2
Count model – a model parameter is given for each independent variable with a confidence interval (in brackets) and a p value. Asterisks indicate statistically sig­
nificant effects (p < 0.05). Deadwood type – L – lying, S – standing; I, II, III – decay stages; 7–14, 15–29, 30–44, 45–59, ≥ 60 – diameter [cm].

Deadwood Constant Stand age [years] Stand volume [m3 Tree density Terrain National parks and
type ha¡1] [trees ha¡1] (upland/mountainous reserves vs. managed
vs. lowland forests) forests

S/I/7–14 0.0399 (− 0.0625; -0.0014 (− 0.0024; 0.0001 (− 0.0002; 0.0003 (0.0002; 0.1111 (0.0719; 0.1503). 0.1969 (0.1158; 0.278).
0.1423). p < 0.001 * − 0.0004). p = 0.007 * 0.0004). p = 0.446 0.0004). p < 0.001 * p < 0.001 * p < 0.001 *
S/I/15–29 0.8667 (0.416; -0.006 (− 0.0102; 0.0014 (0.0002; -0.0003 (− 0.0006; 0). 0.244 (0.0836; 0.4044). 1.0153 (0.6025; 1.4282).
1.3173). p < 0.001 * − 0.0017). p = 0.006 * 0.0026). p = 0.023 * p = 0.078 p = 0.003 * p < 0.001 *
S/I/30–44 0.4592 (− 0.3427; -0.001 (− 0.0111; 0.0023 (− 0.0007; 0 (− 0.0009; 0.0009). 0.4128 (0.0362; 0.7894). 1.4053 (0.4724; 2.3382).
1.2611). p < 0.001 * 0.0092). p = 0.85 0.0054). p = 0.135 p = 0.984 p = 0.033 * p = 0.004 *
S/I/45–59 0.4475 (− 0.8543; 0.0218 (− 0.0191; -0.0014 (− 0.0116; 0.0007 (− 0.0014; 0.199 (− 0.8658; 1.2638). 1.242 (− 0.9126; 3.3966).
1.7492). p = 0.035 * 0.0626). p = 0.302 0.0089). p = 0.795 0.0028). p = 0.532 p = 0.716 p = 0.265
S/I/> 60 14.293 (− 9.1605; 0.0366 (− 0.0924; -0.0255 (− 0.0631; -0.0077 (− 0.034; -1.005 (− 6.0404; 0.7642 (− 2.3232;
37.7464). p = 0.237 0.1656). p = 0.594 0.012). p = 0.22 0.0187). p = 0.583 4.0305). p = 0.706 3.8515). p = 0.641
S/II/7–14 0.1708 (0.058; -0.0039 (− 0.0052; 0.0007 (0.0003; 0.0002 (0.0001; 0.0964 (0.044; 0.1488). 0.5605 (0.4303; 0.6907).
0.2835). p < 0.001 * − 0.0027). p < 0.001 * 0.001). p < 0.001 * 0.0003). p < 0.001 * p < 0.001 * p < 0.001 *
S/II/15–29 1.2686 (0.7844; -0.0121 (− 0.0164; 0.0025 (0.0012; -0.0007 (− 0.001; 0.2674 (0.0742; 0.4606). 1.8853 (1.2652; 2.5054).
1.7527). p < 0.001 * − 0.0077). p < 0.001 * 0.0038). p < 0.001 * − 0.0004). p < 0.001 * p = 0.007 * p < 0.001 *
S/II/30–44 1.2364 (0.3347; 0.0035 (− 0.0085; -0.0002 (− 0.0034; -0.0005 (− 0.0014; 0.0535 (− 0.2946; 2.5329 (1.3414; 3.7245).
2.1382). p < 0.001 * 0.0155). p = 0.567 0.0029). p = 0.882 0.0004). p = 0.254 0.4015). p = 0.764 p < 0.001 *
S/II/45–59 7.8216 (3.5108; -0.0074 (− 0.0307; -0.0055 (− 0.0126; -0.0056 (− 0.0092; -0.5158 (− 1.4157; 1.4347 (0.3028; 2.5666).
12.1324). p < 0.001 * 0.0159). p = 0.538 0.0015). p = 0.129 − 0.0021). p = 0.003 * 0.3841). p = 0.265 p = 0.016 *
S/II/> 60 0.9244 (− 1.399; 0.0272 (− 0.0156; -0.002 (− 0.0149; 0.0009 (− 0.0029; -0.4269 (− 2.3303; -0.508 (− 2.3065; 1.2905).
3.2478). p = 0.119 0.0699). p = 0.226 0.0109). p = 0.763 0.0048). p = 0.637 1.4765). p = 0.665 p = 0.585
S/III/7–14 0.1735 (0.1118; -0.0012 (− 0.0018; 0.0001 (− 0.0001; -0.0001 (− 0.0001; 0). 0.0157 (− 0.0061; 0.1325 (0.0906; 0.1744).
0.2352). p < 0.001 * − 0.0006). p < 0.001 * 0.0002). p = 0.404 p = 0.014 * 0.0375). p = 0.16 p < 0.001 *
S/III/15–29 0.5249 (0.1911; -0.0033 (− 0.0065; 0.0004 (− 0.0005; -0.0002 (− 0.0005; 0). 0.2064 (0.0967; 0.3161). 0.605 (0.3621; 0.8478).
0.8586). p < 0.001 * − 0.0002). p = 0.037 * 0.0013). p = 0.372 p = 0.078 p < 0.001 * p < 0.001 *
S/III/30–44 0.3183 (− 0.261; 0.0033 (− 0.0012; -0.0001 (− 0.0013; -0.0002 (− 0.0007; 0.0134 (− 0.1159; 0.5806 (0.3284; 0.8329).
0.8976). p < 0.001 * 0.0078). p = 0.149 0.001). p = 0.806 0.0003). p = 0.539 0.1427). p = 0.839 p < 0.001 *
S/III/45–59 -0.208 (− 1.0962; 0.0033 (− 0.0061; 0.0007 (− 0.0017; 0.0002 (− 0.0006; 0.1598 (− 0.1397; 0.5046 (0.0911; 0.9181).
0.6803). p = 0.087 0.0128). p = 0.494 0.0032). p = 0.559 0.001). p = 0.621 0.4593). p = 0.301 p = 0.021 *
S/III/≥ 60 3.2198 (− 0.9211; 0.0112 (− 0.0385; -0.0039 (− 0.0163; -0.0029 (− 0.0072; 0.4174 (− 0.9493; 1.3281 (− 0.512; 3.1683).
7.3606). p = 0.059 0.0609). p = 0.663 0.0085). p = 0.546 0.0013). p = 0.187 1.7841). p = 0.556 p = 0.172
L/I/7–14 0.1044 (0.0399; -0.0005 (− 0.0012; 0 (− 0.0002; 0.0002). 0 (0; 0.0001). 0.1049 (0.0758; 0.1341). 0.0776 (0.0237; 0.1315).
0.169). p < 0.001 * 0.0003). p = 0.22 p = 0.78 p = 0.073 p < 0.001 * p = 0.005 *
L/I/15–29 0.3892 (0.0305; 0.0025 (− 0.0017; -0.0006 (− 0.0017; 0 (− 0.0003; 0.0002). 0.1925 (0.0481; 0.3369). 0.515 (0.1693; 0.8607).
0.7479). p < 0.001 * 0.0068). p = 0.241 0.0006). p = 0.337 p = 0.739 p = 0.009 * p = 0.004 *
L/I/30–44 0.5396 (− 0.1651; 0.0086 (− 0.0002; -0.0016 (− 0.0041; -0.0002 (− 0.0008; 0.3197 (0.0305; 0.609). 0.9731 (0.2838; 1.6625).
1.2443). p < 0.001 * 0.0174). p = 0.057 0.0008). p = 0.184 0.0004). p = 0.535 p = 0.032 * p = 0.006 *
L/I/45–59 -0.3446 (− 1.7557; 0.0116 (− 0.0039; -0.0017 (− 0.0056; 0.0007 (− 0.001; 0.5016 (0.054; 0.9493). 0.6271 (− 0.0469;
1.0664). p = 0.369 0.0272). p = 0.151 0.0021). p = 0.374 0.0024). p = 0.44 p = 0.035 * 1.3011). p = 0.077
L/I/> 60 -6.6006 (− 24.5069; 0.0096 (− 0.0583; 0.0104 (− 0.0247; 0.0038 (− 0.0065; 0.8124 (− 1.5139; —
11.3057). p = 0.583 0.0775). p = 0.8 0.0454). p = 0.603 0.014). p = 0.526 3.1387). p = 0.543
L/II/7–14 0.1889 (0.0911; -0.0025 (− 0.0036; 0.0005 (0.0002; 0 (0; 0.0001). 0.1095 (0.0645; 0.1544). 0.5651 (0.452; 0.6782).
0.2867). p < 0.001 * − 0.0014). p < 0.001 * 0.0008). p = 0.001 * p = 0.475 p < 0.001 * p < 0.001 *
L/II/15–29 0.346 (0.0838; -0.0034 (− 0.0067; 0.0015 (0.0007; -0.0003 (− 0.0005; 0.3096 (0.1835; 0.4358). 1.9876 (1.5275; 2.4477).
0.6083). p < 0.001 * − 0.0002). p = 0.036 * 0.0024). p < 0.001 * − 0.0001). p = 0.005 * p < 0.001 * p < 0.001 *
L/II/30–44 0.3469 (− 0.0565; -0.0012 (− 0.0063; 0.0013 (− 0.0001; -0.0004 (− 0.0008; 0). 0.1357 (− 0.0219; 1.747 (1.2324; 2.2616).
0.7504). p < 0.001 * 0.0039). p = 0.643 0.0027). p = 0.064 p = 0.037 * 0.2934). p = 0.093 p < 0.001 *
L/II/45–59 0.7799 (− 0.3288; -0.0014 (− 0.0221; 0.0004 (− 0.0048; -0.0003 (− 0.0016; 0.2437 (− 0.3089; 0.9792 (0.0139; 1.9445).
1.8886). p = 0.003 * 0.0193). p = 0.895 0.0055). p = 0.889 0.001). p = 0.635 0.7963). p = 0.391 p = 0.051
L/II/> 60 1.6277 (− 1.787; -0.0183 (− 0.1037; 0.009 (− 0.0178; -0.001 (− 0.0064; -2.2426 (− 4.5361; -0.3718 (− 2.2707;
5.0425). p = 0.157 0.0671). p = 0.682 0.0358). p = 0.524 0.0044). p = 0.719 0.0509). p = 0.079 1.5271). p = 0.708
L/III/7–14 0.3764 (0.269; -0.0025 (− 0.0037; 0.0003 (− 0.0001; -0.0001 (− 0.0002; 0.1174 (0.0706; 0.1642). 0.7059 (0.5819; 0.8299).
0.4838). p < 0.001 * − 0.0013). p < 0.001 * 0.0006). p = 0.117 − 0.0001). p < 0.001 * p < 0.001 * p < 0.001 *
L/III/15–29 0.43 (0.1144; 0.7457). -0.0017 (− 0.0051; 0.001 (0.0001; -0.0003 (− 0.0005; 0.6864 (0.5251; 0.8477). 2.3949 (1.8226; 2.9673).
p < 0.001 * 0.0018). p = 0.347 0.002). p = 0.035 * − 0.0001). p = 0.007 * p < 0.001 * p < 0.001 *
L/III/30–44 0.0862 (− 0.4022; 0.0054 (− 0.0007; -0.0001 (− 0.0016; 0 (− 0.0004; 0.0004). 0.5228 (0.3152; 0.7304). 2.0119 (1.3485; 2.6754).
0.5747). p < 0.001 * 0.0114). p = 0.084 0.0015). p = 0.944 p = 0.976 p < 0.001 * p < 0.001 *
L/III/45–59 0.4289 (− 0.735; -0.0034 (− 0.0149; 0.0016 (− 0.0013; -0.0003 (− 0.0012; 0.3983 (0.0097; 0.7869). 1.7787 (0.7211; 2.8363).
1.5928). p = 0.018 * 0.008). p = 0.559 0.0046). p = 0.281 0.0006). p = 0.548 p = 0.047 * p = 0.001 *
L/III/≥ 60 0.2197 (− 0.8767; -0.013 (− 0.0524; 0.0044 (− 0.0064; 0.0011 (− 0.0013; 0.1343 (− 0.9174; 2.8506 (0.0699; 5.6312).
1.316). p = 0.036 * 0.0263). p = 0.52 0.0152). p = 0.431 0.0036). p = 0.374 1.1861). p = 0.804 p = 0.053

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