Final Thesis
Evaluation of artificial habitats for saproxylic oak
invertebrates: Effects of substrate, composition
and distance from dispersal source
Anna Larsson
LiU-IFM-Ex--08/1893--SE
Rapporttyp Report category
Licentiatavhandling
x Examensarbete
C-uppsats x D-uppsats
Övrig rapport
x_ _45 hp_______________
Språk Language
Svenska/Swedish
x Engelska/English
_ ________________
Titel/Title Utvärdering av artificiella miljöer för saproxyliska ekevertebrater: effekter på substrat,
sammansättning och avstånd från spridningskällor
Evaluation of artificial habitats for saproxylic oak invertebrates: Effects of substrate,
composition and distance from dispersal source
Författare/Author Anna Larsson
Sammanfattning/Abstract
Saproxylic species living in old hollow trees have low dispersal rate. Many of the
species are threatened since their micro habitats are rare. To prevent some of these
species from going extinct their habitats have to have the right management. In some
areas artificial environment could be a solution. The aim of this study was to investigate
if the insects that are dependent on tree cavities with wood mould would colonize an
artificially created habitat: large wooden boxes filled with artificial wood mould placed
on tree trunks. The boxes were filled with substrates like oak saw dust, oak leaves, dead
hens, hen excrements, medicago (Medicago falcata flour) or potatoes. Over three years,
136 species and 10 380 specimens were caught in 47 boxes. The groups classified as
specialists were in general statistically significant more often than groups classified as
generalists. Dead hen was the substrate with the highest number of species, although
differences were small.
In conclusion, a large number of species, including red listed ones and saproxylic
specialists used the boxes. A dead hen in the box gave some extra species and 1800
meters was too long for some of the species to disperse. Hence, the prospects for using
artificial environments are good especially to reduce habitat availability gaps in time and
space.
ISBN
LiU-IFM-Ex--08/1893--SE
____________________________________
_________________
ISRN LiU-Biol-Ex-614
____________________________________
Handledare: Professor Per Milberg
Nyckelord/Keyword
artificial environment, hollow oak, Quercus robur, saproxylic beetles, Sweden, wood
mould boxes
Datum/Date
10/01/2008
URL för elektronisk version
Avdelning, Institution
Division, Department
Avdelningen för ekologi
Institutionen för fysik, kemi och biologi
Contents
1 Abstract ...................................................................................................... 1
2 Introduction ................................................................................................ 1
3 Materials and methods ............................................................................... 3
3.1 Field study ............................................................................................ 3
3.1.1 Study sites ...................................................................................... 3
3.1.2 Traps and boxes ............................................................................. 4
3.1.5 Animals identification ................................................................... 6
3.2 Data analyses ....................................................................................... 6
4 Results ........................................................................................................ 7
4.1 Distance effect ..................................................................................... 9
4.1.1 Year 1 and 2 .................................................................................. 9
4.1.2 Year 3 ............................................................................................ 9
4.2 Substrate effect .................................................................................... 9
4.2.1 Year 1 and 2 .................................................................................. 9
4.2.2 Year 3 .......................................................................................... 10
4.3 Time effect ......................................................................................... 10
4.4 Assemblage ........................................................................................ 10
5 Discussion ................................................................................................ 12
6 Acknowledgements .................................................................................. 14
7 References ................................................................................................ 15
Appendix ..................................................................................................... 19
Appendix A .............................................................................................. 19
Appendix B .............................................................................................. 22
1
1 Abstract
Saproxylic species living in old hollow trees have low dispersal rate. Many
of the species are threatened since their micro habitats are rare. To prevent
some of these species from going extinct their habitats have to have the
right management. In some areas artificial environment could be a solution.
The aim of this study was to investigate if the insects that are dependent on
tree cavities with wood mould would colonize an artificially created
habitat: large wooden boxes filled with artificial wood mould placed on
tree trunks. The boxes were filled with substrates like oak saw dust, oak
leaves, dead hens, hen excrements, medicago (Medicago falcata flour) or
potatoes. Over three years, 136 species and 10 380 specimens were caught
in 47 boxes. The groups classified as specialists were in general statistically
significant more often than groups classified as generalists. Dead hen was
the substrate with the highest number of species, although differences were
small.
In conclusion, a large number of species, including red listed ones and
saproxylic specialists used the boxes. A dead hen in the box gave some
extra species and 1800 meters was too long for some of the species to
disperse. Hence, the prospects for using artificial environments are good
especially to reduce habitat availability gaps in time and space.
Keywords: artificial environment, hollow oak, Quercus robur, saproxylic
beetles, Sweden, wood mould boxes
2 Introduction
Old oaks (Quercus robur, Q. petraea) sustain a diverse fauna of saproxylic
beetles (Palm 1959). The definition of saproxylic species is species of
invertebrates that are dependent, during some part of their life cycle, upon
the dead wood of moribund or dead trees (standing or fallen), or upon
wood-inhabiting fungi, or upon the presence of other saproxylics (Speight
1989b). At least 6000-7000 species in Sweden are saproxylic (Dahlberg &
Stokland 2004). The Swedish fauna includes approximately 1000
saproxylic beetle species (Esseen et al. 1997), many of which are red listed
(Gärdenfors 2005).
In the trunks of old oaks, hollows with wood mould are often formed.
Wood mould consists of loose wood colonized by fungi, often with remains
from insects and animal nests. Oaks with wood mould hollows harbour a
specialized fauna, mostly consisting of beetles and flies (Dajoz 1980,
Ranius 2002a).
2
Some species are totally dependent on a temporal continuity of oaks
for their survival (Hedin 1999). They have been shown to have a low
dispersal rate and range in comparison with saproxylic beetles in other
microhabitats (Ranius & Hedin 2001). Many species associated with old
oaks in Europe are threatened because of the decreasing habitat (Ranius &
Jansson 2002). One well-studied example is Osmoderma eremita (Luce
1996, Ranius & Hedin 2001, Ranius 2002b), which is also one of the
species with the highest priority in the European Union’s Habitat Directive
(Luce 1996).
To be able to have large old oaks in an open landscape, the habitat
needs disturbance e.g. grazing (Ranius 2006) or fire (Glasgow & Matlack
2007). Only a small part of such habitats remain today. If the old oaks have
strong competition from younger trees that shade the oaks, they will die
due to lack of light (Read 2000). The large gap of oaks is one big reason
that can lead to local extinction of species that are oak specialists. It is also
possible that an extinction dept exist because of historical or recent changes
in the abundance of old oaks (Ranius 2002c, Ranius & Kindvall 2006).
To try to understand population dynamics and estimate the extinction
risks it is important to know as much as possible of species’ dispersal rate
and range (Ranius 2006). The dispersal is necessary for gene flow (Slatkin
1987) and for colonization of unoccupied patches (Hanski et al. 1994).
Dispersal is also important to recolonization of patches with previous
extinctions of local populations (Brown & Kodrick-Brown 1977). Dispersal
is an important factor to cope with if the environment changes, for example
habitat loss and fragmentation (Thomas 2000). The knowledge of the
dispersal capacity for the saproxylic invertebrates living on old hollow oaks
is poor but studies have shown that one of the most stable dead-wood
microhabitats is likely the interior of a hollow tree and beetles inhabiting
this environment seem to have low dispersal rates (Nilsson & Baranowski
1997). Among saproxylic insects, bark beetles and fire dependent species
are well known for being good dispersers (Forsse & Solbreck 1985, Wikars
1997). There are sites with unique fauna where the age structure will create
lack of hollow oaks in the future or where the distance to the next species-
rich patch is long. In this situation, one possible solution could be to create
artificial environments to fill the spatial and temporal habitat gap.
The aim of this study was to investigate if the insects that are
dependent on tree cavities with wood mould will colonize the created
habitat.
3
The hypotheses evaluated were:
Distance does affect a) species living in the cavity of hollow trees, b)
species of conservation interest, c) red listed saproxylic species and d)
obligate saproxylic beetle species. In contrast, we expected no apparent
distance effect for e) species living in bird nests or nests of social insects
(ants and wasps) and f) fungi. A distance effect is expected because these
species are intimately confined to their substrates.
I expected that the substrate in the wood mould boxes affects a)
species living in the cavity of hollow trees, e) species living in bird nests or
nests of social insects (ants and wasps) and g) richness. A substrate effect is
expected since some species are dependent on specific substrates.
I also expected to detect a time effect for a) species living in the cavity
of hollow trees and g) richness. I think the substrates will be affected by
time, for that reason species may be attracted in different periods and also
the species composition will be affected.
3 Materials and methods
3.1 Field study
3.1.1 Study sites
The different study sites, Brokind, Bjärka Säby and Grebo, are situated
about 15-20 kilometres south to south-east of Linköping, Sweden (Figure
1).
4
Figure 1. Locations of the three study sites, south to south-east of Linköping, in the county of Östergötland, Sweden.
The specific study sites were selected because they are known to harbour a
rich saproxylic invertebrate fauna living in old oaks (Ranius & Jansson
2002). Another reason for choosing these sites was the lack of hollow oaks
in one or more directions from the core area of the sites. The study sites
had between 50-100 hollow oaks each.
3.1.2 Traps and boxes
Both pitfall traps and eclector traps were used to collect the beetles. To try
to resemble the conditions in big hollow oaks, for example in temperature
and moist, large wooden boxes were used. There were 47 boxes used in the
study. The boxes were made of oak board (2.5 cm in the walls and 5 cm in
the bottom). The size was about 0.70 x 0.30 x 0.30 meters and a volume of
about 60 liters. They were kept together with brass screws. The bottom
inside of each box was covered with 50 mm clay, formed as a bowl, to keep
the moisture. The boxes looked like large bird nests and were filled with
70 % artificial wood mould components. The artificial wood mould
consisted of oak saw dust, oak leaves, lucerne flour (Medicago falcata),
hay and water. The boxes were also filled with different substrates like
potatoes, lucerne flour, hen’s excrements or dead hens (Table 1). The
potatoes were used to mimic a moist environment and lucerne flour was put
into some boxes to create an extra high protein content. Hen’s excrement
5
and dead hen were used to resemble the circumstances in hollow trees with
bird nests.
Table 1. The total number of boxes per distance and substrate.
Distances (meter)
Total number of
boxes No. of boxes with hen’s excrement
No. of boxes with potato
No. of boxes with medicago
No. of boxes with dead hen
0 12 3 3 3 3
100 8 2 2 2 2
200 3 1 1 1 0
300 4 1 1 1 1
400 4 1 1 1 1
600 8 2 2 2 2
1800 8 2 2 2 2
The boxes were placed about 4 meters up on the trunk on the north side or
in the shade on oaks. They stayed in position with metallic band around the
tree. The roof and one side of the box could be opened but behind the door
at the side there was a transparent plastic window. There were small holes
and some milled notches in the roof of the box to lead some rain water in
(Figure 2).
Figure 2. A schematic drawing of a box with notches and holes on the roof to let some rain water in. The clay was formed as a bowl in the bottom to help retain moisture.
The boxes were placed in three different core areas. Each core area was
divided into a central area and two or three smaller areas in different
directions and distances from the central area. The distances were 0 meter,
100 meters, 200 meters, 300 meters, 400 meters, 600 meters and 1800
meters. The boxes stayed out for four seasons. The first season the boxes
were opened, the animals could reach the inside of the boxes through holes
(Figure 3), without disturbance. The second and third seasons they were
opened and had pitfall traps. The pitfall traps were plastic jars with a top
Milled notches
Holes
Clay
6
diameter of 70 mm, placed with the opening level within the wood mould
in the hollows. The pitfall traps were filled with 70 % of conservation fluid.
The conservation fluid consisted of one half of glycol and one half of water
and a small part of detergent to eliminate surface tension. The pitfall traps
were placed in the boxes one week at a time, about three times each year,
between May to August. During the fourth and last season the boxes were
closed (Figure 3), covered with dark plaid and hatching colonizers were
caught with a kind of eclector trap. On one side of the trap there was a hole,
80 mm in diameter, where a white plastic bottle was placed. This was the
only place where light came into the trap. Beetles emerging from the trap
were caught in the bottle. The bottle was kept in position with a metallic
piece. The plastic bottle was filled with about 25% of conservation fluid
and the bottles were changed about once a month from May to September.
The aim was to catch all the hatching insects who were attracted to the light
from the bottle.
Figure 3. A schematic sketch of which years the boxes were opened (white dots) and when the boxes were closed with eclector traps (black dots).
3.1.5 Animals identification
The traps were managed by Arne Ekström during 2002-2003 who also
identified the caught invertebrates. During 2003-2007 the caught
invertebrates were taken to laboratory where they were sorted and
identified by Nicklas Jansson and Anna Larsson in 2007. Some groups
were sent to other specialists for identification. Staphylinidae were
identified by Stig Lundberg, Cryptophagidae by Rickard Andersson,
Pseudoscorpions by Stanislav Snäll and Syrphidae and Tipulidae by Hans
Bartsch.
3.2 Data analyses
Statistica 7 (Statsoft Inc 2004) Generalized Linear/Nonlinear Model (GLZ)
with Poisson distribution and log link function was used to evaluate the
effects of substrates, years and distances. As a number of tests were
7
conducted, the Holm correction for multiple testing was applied. The full
species assemblage recorded in the boxes was analysed in Canonical
Corresponding Analyses (CCA) with the software CANOCO 4.5 with log
transformation (ter Braak & Smilauer 2002). Each of the experimental
factors was evaluated using the others as covariables, hence conducting a
number of partial CCA (pCCA). For the data from year 1-2 (i) time, (ii)
substrate and (iii) distance were evaluated; for year 3 only the two latter.
The statistical significance of the species composition was evaluated in
Monte Carlo tests using 9999 permutations (using permutation blocks
defined by the covariables). A partial Detrended Correspondence Analysis
(pDCA) was made with Canoco 4.5 to investigate if the factors that we
could not control (vespa nest, ant nest, cardinal points, sun exposure or the
different areas) had any impact on the results. A comparison was made
with species found in the studied areas in another study using window traps
and pitfall traps in 1994 (Ranius & Jansson 2002) and the species recorded
in the present study. In this case branch-classed species were excluded as
this group was of low interest in the present study.
4 Results
In total 136 species (Table 2) and 10 380 specimen were found (Appendix
B, Table 1). The most species rich box had 37 species. From the Swedish
Red list (Gärdenfors 2005) there was a maximum four red listed species in
a box. The mean value of species per box was 17 (SD 7.9). The mean
number of specimens per box was 430 (SD 471.7). The Simpson diversity
index (Magurran 2004) mean value was 0.92 (SD 0.068). A ranking
between the substrates showed that the dead hen gave the highest number
of specimens and species (Table 3). The pDCA did not indicate that the
factors we could not control had any impact on the results of the study
(data not shown).
8
Table 2. A summary of how many species of each category that were found in the study (classifications of species is found in Appendix B, Table 1).
Number of Proportion
Species year 1, 2 and 3 136
Facultative saproxylic beetle species 51
Obligate saproxylic beetle species 53
Species of conservation interest 28
Swedish Red list 2005 species 14
Pseudoscorpion species 5
Hover fly species 4
Crane fly species 2
Ant species 5
Dry wood species 4
Tree hollow species 17
Nest of birds, ants or wasps species 19
Rotting wood species 19
Species compared to Ranius & Jansson 2002 57 45.6 %
Species of conservation interest compared to Ranius & Jansson 2002 20 39.6 %
Red list 2005 species compared to Ranius & Jansson 2002 9 31.0 %
Table 3. A summary of the significant test outcomes for the substrate variable (c.f. Appendix A, Table 1). For each significant test, substrates were ranked from the one with the highest number of specimens or species (3) to the lowest (0).
Categories
Specimen/ Species
level Year Holms
correction Hen's
excrement Dead hen Potatoes Medicago
Tree hollow Specimen 1&2 *** 3 1 0 2 Nest of birds, ants
or wasps Specimen 1&2 *** 0 3 2 1
Nest of birds, ants or wasps Species 1&2 *** 1 3 0 2 Obligate
saproxylic beetles Specimen 1&2 *** 0 3 1.5 1.5 Species of
conservation interest Specimen 1&2 *** 0 3 1 2 Swedish Red list
2005 Specimen 1&2 *** 0 3 1 2 Tree hollow Specimen 3 *** 2 3 0 1
Nest of birds, ants or wasps Specimen 3 *** 2 3 1 0
Facultative saproxylic beetles Specimen 3 *** 1 3 0 2
Obligate saproxylic beetles Specimen 3 *** 2 3 1 0
Rotting wood Specimen 1&2 * 2 0 1 3
Crane flies Specimen 3 * 2 0 1 3 Species of
conservation interest Specimen 3 * 2 3 0 1
Proportion 22,67% 41,33% 10,67% 25,33%
9
4.1 Distance effect
Compared with the expected outcomes, there were conflicting results when
testing for the distance effect. The groups “tree hollow”, “obligate
saproxylic beetle species” and “Swedish Red list 2005” were expected to
exhibit a decrease of specimens/species with increasing distance.
When analysing the full species assemblage, there was a highly
significance distance effect (p=0.0001 for both year 1 & 2 and year 3
(Table 4)). The species most strongly affected by distance, positively or
negatively, are found in Table 5.
4.1.1 Year 1 and 2
The number of specimens of “tree hollow” (decreasing with increasing
distance) and “Swedish Red list 2005” (increasing with increasing distance)
were highly significant (Appendix A, Table 1). “Obligate saproxylic beetle
species” at specimen level (increasing with increasing distance) were
statistical significant and also “tree hollow” at species level (decreasing
with increasing distance) (Appendix A, Table 1).
4.1.2 Year 3
Statistically significant results on number of specimen and species were
found in the groups “nest of birds, ants or wasps” (increasing with
increasing distance), “obligate saproxylic beetle species” (increasing with
increasing distance), “species of conservation interest” (increasing with
increasing distance) and “Swedish Red list 2005” (decreasing with
increasing distance) (Appendix A, Table 1).
4.2 Substrate effect
The groups “tree hollow”, “nest of birds, ants or wasps” and “obligate
saproxylic beetle species” were expected to be highly statistically
significant, but the results of GLZ were not easily interpret.
4.2.1 Year 1 and 2
At specimen level “tree hollow”, “nest of birds, ants or wasps”, “obligate
saproxylic beetle species”, “species of conservation interest” and “Swedish
Red list 2005” were highly significant. Also “nest of birds, ants or wasps”
at species level were highly significant. “Rotting wood” at specimen level
was statistically significant (Appendix A, Table 1).
Figure 4 describes the relationship between the different substrates and
the species assemblage, and selections of the species are indicated. Some
species e.g. Trox scaber, Nemadus colonides, Gnathoncus buyssoni and
Quedius brevicornis were associated with the dead hen substrate. Some
10
species e.g. Ptinus fur, Atetha nigricornis and Dendrophilus corticalis were
in the centre of the graph and hence seemed unaffected by substrates.
4.2.2 Year 3
At specimen level “facultative saproxylic beetle species”, “tree hollow”,
“nest of birds, ants or wasps” and “obligate saproxylic beetle species” were
highly significant. “Crane flies” and “species of conservation interest” at
specimen level were statistical significant (Appendix A, Table 1).
4.3 Time effect
“Nest of birds, ants or wasps” and “tree hollow” was expected to be highly
significant with an increase of specimens/species with increasing year, but
the GLZ results showed the opposite.
“Tree hollow” and “rotting wood” at specimen level were statistically
significant with an increase of specimens/species. “Facultative saproxylic
beetle species”, “nest of birds, ants or wasps”, “obligate saproxylic beetle
species” and “Swedish Red list 2005” at specimen level were statistically
significant with a decrease of specimen/species (Appendix A, Table 1).
4.4 Assemblage
It was only substrate in year three that were not statistical significant in the
Monte Carlo tests (Table 4).
Table 4. Summary of Monte Carlo tests, 9999 permutations under reduced model.
Year 1&2 Year 3
p-value p-value
Distances 0.0001 Substrate and year
as co-variables 0.0001 Substrate as co-variables
Substrates 0.0127 Year as co-variables 0.5404 Distance as co-variables
Year 0.0006 Substrate as co-variables
11
Table 5. Ranking of species according to their association with increasing (+) or decreasing (-) distance from the core area in two pCCAs, sorting according to mean ordination score. The species were chosen from two criteria, at least ten individuals and at least six (Year 1 & 2) or three (Year 3) in frequency.
Year 1 & 2 Year 3
Sp
ec
ies
Ra
nk
ing
EIG
. 0
.33
92
Ab
un
da
nce
Fre
qu
en
cy
Ra
nk
ing
EIG
. 0
.41
97
Ab
un
da
nce
Fre
qu
en
cy
Pti fur 2 -1.66 464 23 1 -2.09 109 16
Myc lin 3 -1.45 21 7
Pri ate 1 -1.83 29 16 3 -1.02 23 8
Lio mar 6 -1.17 10 9
Pti sub 4 -1.43 20 9 4 -0.77 20 12
Dic bim 7 -0.97 15 13 2 -1.23 13 7
Den cor 5 -1.19 29 13 5 -0.55 25 4
Vel dil 9 -0.63 38 8
Cor ser 8 -0.78 102 23 10 -0.27 11 5
Cry sca 13 0.20 72 20 7 -0.43 27 12
Phi sub 15 0.21 140 23 8 -0.41 17 6
Ate nig 16 0.30 600 48 9 -0.35 192 20
Tro sca 12 0.19 57 21 12 0.03 48 10
Gna buy 18 0.74 44 19 6 -0.51 11 3
Eup fau 15 0.14 25 12
Que bre 14 0.21 161 27 14 0.12 14 3
Scr fus 16 0.25 42 10
Nem col 11 -0.37 45 20 20 0.99 20 10
Hap vil 17 0.64 227 28 13 0.03 38 6
Tha hos 20 1.26 54 10 11 -0.23 17 4
Eup kar 17 0.58 127 20
Hap mel 10 -0.50 12 7 23 1.71 37 4
Cry con 18 0.68 11 3
Cte pec 19 0.75 64 30 19 0.87 48 14
Hap nig 21 1.27 103 4
Cry mic 22 1.59 62 4
Din pan 21 1.67 12 7
Sep tes 22 1.69 15 8
Bra lap 23 1.82 14 9
Cer his 24 2.13 33 16
Eup nan 24 2.91 17 5
12
Figure 4. A CCA (Canoco 4.5) of year 1 and 2 that shows the association of some species and the different substrates. The smallest triangles ( ) correspond to <20 individuals, next size ( ) correspond to 21-100 individuals,
next size ( ) correspond to 101-300 individuals and the larges triangles ( ) correspond to >300 individuals. The white triangles show that species are found in <10 boxes, grey triangles 11-30 boxes and black triangles in 31-94 boxes.
5 Discussion
Forest management in northern Europe has been very intensive during the
last decades. Many species associated with dead trees and decaying wood
have decreased (Essen et al. 1992, Haila 1994, Siitonen & Martikainen
1994) and a large number of these species are now threatened or vulnerable
(Ehnström et al. 1993). A lot of the species that are dependent on old, large
hollow trees have survived in small remnant woodlands with old trees,
often in the agricultural landscape (Speight 1989a, Warren & Key 1989).
As expected the “tree hollow” species and “Swedish Red list 2005”
exhibited a highly significant distance effect. Species in the group “tree
hollow” were decreasing with increasing distance and one explanation is
because they have low mean dispersal rates (Ranius & Hedin 2001).
-1.5 2.5
-2.0
2.0
Gna buy
Den cor
Nem col
Vel dil
Tha hos
Que bre
Hap vil
Ate nig
Tro sca
Pti furPti sub
Cry sca
Cer his
Cor ser
Pri ate
Cte pec
Dic bim
EXCREMENT
POTATO
DEAD HEN
MEDICAGO
EIG AX1 0.1987
13
The hollow is a stable environment, compared with the life cycle of
invertebrates, and lots of animals have no need of moving (Ranius & Hedin
2001) and historically it has been better to stay in a stable and long-lived
environment than looking for a new one and risking to fail.
The boxes were used by the species as we expected, especially the
groups “tree hollow”, “nest of birds, ants or wasps” and “Swedish Red list
2005”. One reason could be that these groups are more specific to their
ecological niches than the others. In one group, fungi, too few
specimens/species were found and no calculations could be made.
The conversion of natural forest has resulted in increased distances
between clusters of patches, which suggest that limited dispersal capacity
may be a problem for threatened species (Warren & Key 1991, Haila et al.
1994). The distances used in this study were chosen from experiences in
earlier studies (Ranius & Hedin 2001). The thought was to have distances
where the species have their maximum range. The obligate saproxylic
beetle species are increasing with increasing distance. That could be
because they are generalists and then have other tree species as their main
substrate. Another possibility is that they have a source in the surrounding
that was missed when choosing the locations.
The substrates were selected to resemble environmental conditions
inside the hollow trees. The hen’s excrement was supposed to resemble
excrement that occurs in the nature and the medicago was to keep a high
protein content in the specific boxes. The hen’s excrement and the
medicago substrates had about the same proportion of animals and the
potato substrate had the lowest proportion when calculating from the mean
values. Probably no animals were attracted to the potato itself, but maybe
to the moisture the potato retained.
It comes as no surprise that Trox scaber, Nemadus colonides, Quedius
brevicornis and Gnathoncus buyssoni are situated near the dead hen
substrate in the multivariate analysis (Figure 4) as it is well known that
these species are attracted to carrions in hollow trees. The idea with placing
a dead hen in a box was to resemble dead birds, like dead nestlings or
carrions from owls or jackdaws, in hollows in the forest. This substrate
resulted in the highest specimen numbers. One reason why e.g.
Dendrophilus corticalis and Quedius brevicornis have a position near the
excrement substrate in the CCA is that they live as predators of the diptera
larvae living in wood mould and hen excrements might have increased their
number in these boxes.
The species categorized as “dry wood” are probably primarily
interested of the oak wood in the boxes, not the substrates itself.
14
Most of the substrates used in this study are likely to lose their
attraction over time. The species that are associated with nests are also
attracted of dead birds. After the first year the dead hen has broken down
and its attraction was much less than the first year. Numbers of specimens
from the group “nest of birds, ants or wasps animals” decreased with time.
“Tree hollow” specimens were increasing with time. That is maybe
because they produce offspring in the unoccupied boxes since they offer a
large amount of suitable environment.
When calculating pDCAs (ter Braak & Smilauer 2002) of the factors
that were not under experimental control, it was difficult to infer any
pattern in the solution. Hence, uncontrolled factors, like vespa nest, ant
nest, areas, cardinal points and sun exposure exerted only a small influence
on the assemblages of species.
If using boxes as a part of management strategy, I would like to stress
that neither size nor material has been explored here. If the boxes are
larger, environmental conditions like moisture and temperature in the box
would be more stabile.
In conclusion, a large number of species, including red listed ones and
saproxylic specialists used the boxes. A dead hen in the box gave some
extra species and 1800 meters seemed too long for some of the species to
disperse. Hence, the prospects for using artificial environments are good
especially in nature conservation.
6 Acknowledgements
I would like to sincerely thank my supervisor professor Per Milberg for all
kind of help with for example statistics, writing and always having time.
Also PhD student Nicklas Jansson has helped me a lot during this study, for
instance with encouragement, species identification and writing. I would
also like to thank Thomas Ranius for interesting comments on the statistics
and all specialists for species identification: Stig Lundberg, Rickard
Andersson, Stanislav Snäll and Hans Bartchs. I would like to thank my
parents Gunilla and Thomas Larsson for helping me during the field work
and together with Johan Carpholm always supporting me and helping me
with computer problems. Last but not least I would like to send big thanks
to everyone who has supported me and had comments during this work.
15
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19
Appendix
Appendix A
Table 1. Total number of species, frequency, Wald statistic, p-value and Holm´s corrected significance level for different groups of insects living close to oaks (Quercus robur, Q. petraea) caught by pitfall traps and eclector traps in Östergötland, Sweden.
Year 1 & 2 Year 3
Ab
un
dan
ce
Fre
qu
en
cy
(nm
ax=
94)
Wald
sta
tisti
c
p-v
alu
e
Inc
reasin
g o
r
de
cre
asin
g
Ho
lm´s
co
rrecti
on
Ab
un
dan
ce
Fre
qu
en
cy
(nm
ax=
47)
Wald
sta
tisti
c
p-v
alu
e
Inc
reasin
g o
r
de
cre
asin
g
Ho
lm´s
co
rrecti
on
Dry wood 1
Specimens 23 9 9 5
sqrtDist 6.17 0.0129 - 1.44 0.231
Substrate 15.0 0.00180 1.71 0.635
Year 3.32 0.0681
No. of species 4 4
sqrtDist 3.069 0.0798 0.957 0.328
Substrate 1.65 0.648 0.928 0.819
Year 0.394 0.530
Tree hollow 1
Specimens 558 42 313 34
sqrtDist 392 <0.0000 - *** 10.6 0.00112 -
Substrate 171 <0.0000 *** 47.8 <0.0000 ***
Year 118 <0.0000 + ***
No. of species 16 16
sqrtDist 14.5 0.000141 - * 1.14 0.286
Substrate 0.452 0.929 1.12 0.773
Year 8.55 0.00346 +
Nest of birds, ants or
wasps 1
Specimens 430 67 238 25
sqrtDist 3.41 0.0649 49.6 <0.0000 + ***
Substrate 155 <0.0000 *** 89.7 <0.0000 ***
Year 31.5 <0.0000 - ***
No. of species 20 19
sqrtDist 3.38 0.0661 0.562 0.453
Substrate 45.6 <0.0000 *** 8.89 0.0308
Year 0.703 0.402
1 Groups defined by Ranius & Jansson (2002)
20
Appendix A, Table 1 continued
Year 1 & 2 Year 3
Ab
un
dan
ce
Fre
qu
en
cy
(nm
ax=
94)
Wald
sta
tisti
c
p-v
alu
e
Inc
reasin
g o
r
de
cre
asin
g
Ho
lm´s
co
rrecti
on
Ab
un
dan
ce
Fre
qu
en
cy
(nm
ax=
47)
Wald
sta
tisti
c
p-v
alu
e
Inc
reasin
g o
r
de
cre
asin
g
Ho
lm´s
co
rrecti
on
Rotting wood
1
Specimens 183 51 82 31
sqrtDist 5.05 0.0246 + 0.242 0.623
Substrate 18.6 0.000331 * 12.3 0.00641
Year 13.8 0.000200 + *
No. of species 19 19
sqrtDist 1.83 0.177 0.236 0.627
Substrate 3.18 0.364 5.41 0.144
Year 1.52 0.217
Pseudo-scorpions
Specimens 23 16 7 5
sqrtDist 8.77 0.00306 + 2.07 0.151
Substrate 8.34 0.0395 3.52 0.318
Year 4.81 0.0283 +
No. of species 4 3
sqrtDist 3.89 0.0486 + 1.59 0.208
Substrate 5.88 0.117 1.35 0.718
Year 2.71 0.100
Hover flies
Specimens 28 15 1 1
sqrtDist 11.1 0.000883 + 0.0316 0.859
Substrate 0.984 0.805 <0.0000 1.00
Year <0.0000 1.00
No. of species 4 1
Crane flies
Specimens 79 36 61 19
sqrtDist 6.02 0.0142 + 2.59 0.107
Substrate 5.45 0.141 17.8 0.000494 *
Year 0.114 0.736
No. of species 2 2 Facultative saproxylic
beetle species
1
Specimens 1749 83 789 44
sqrtDist 10.5 0.00122 + 2.00 0.157
Substrate 15.1 0.00173 131 <0.0000 ***
Year 31.4 <0.0000 - ***
1 Groups defined by Ranius & Jansson (2002)
21
Appendix A, Table 1 continued
Year 1 & 2 Year 3
Ab
un
dan
ce
Fre
qu
en
cy
(nm
ax=
94)
Wald
sta
tisti
c
p-v
alu
e
Inc
reasin
g o
r
de
cre
asin
g
Ho
lm´s
co
rrecti
on
Ab
un
dan
ce
Fre
qu
en
cy
(nm
ax=
47)
Wald
sta
tisti
c
p-v
alu
e
Inc
reasin
g o
r
de
cre
asin
g
Ho
lm´s
co
rrecti
on
Facultative saproxylic
beetle species
1
No. of species 55 54
sqrtDist 3.64 0.0566 0.0698 0.792
Substrate 11.5 0.00939 6.77 0.0795
Year 0.0327 0.856 Obligate
saproxylic beetle
species 1
Specimens 562 78 322 37
sqrtDist 18.3 0.0000191 + ** 42.4 <0.0000 + ***
Substrate 92.7 <0.0000 *** 53.8 <0.0000 ***
Year 32.3 <0.0000 - ***
No. of species 58 58
sqrtDist 2.85 0.0915 0.764 0.382
Substrate 11.6 0.00896 3.54 0.316
Year 0.00469 0.945
Species of conservation
interest 2
Specimens 231 54 214 31
sqrtDist 2.40 0.121 20.7 <0.0000 + ***
Substrate 30.2 <0.0000 *** 17.2 0.000632 *
Year 2.70 0.101
No. of species 23 19
sqrtDist 0.0905 0.763 0.913 0.339
Substrate 3.45 0.328 3.34 0.342
Year 0.274 0.600
Swedish Red list 2005
Specimens 151 34 53 17
sqrtDist 80.6 <0.0000 + *** 14.7 0.000125 - **
Substrate 41.4 <0.0000 *** 12.2 0.00682
Year 45.1 <0.0000 - ***
No. of species 12 9
sqrtDist 0.667 0.414 0.393 0.531
Substrate 4.35 0.226 1.52 0.677
Year 2.94 0.0866
1 Groups defined by Ranius & Jansson (2002)
2 Defined as spp on the Swedish Red list 2000 (Gärdenfors 2000)
22
Appendix B
Table 1. Table of all species, some categories, number of individuals and the frequency.
Sp
ec
ies
Ca
t. N
o.
1
Sp
ec
ies o
f c
on
se
rva
tio
n
inte
res
t 2
Tre
ath
ca
teg
ory
Sw
ed
ish
Re
d l
ist
20
05
En
vir
on
men
t 3
Sa
pro
xy
lic
Fac
ult
ati
ve
=F
Sa
pro
xy
lic
Ob
lig
ate
=O
3
Ye
ar
1
Ye
ar
2
Ye
ar
3
No
. o
f in
div
idu
als
Fre
qu
en
cy
No
. o
f in
div
idu
als
Fre
qu
en
cy
No
. o
f in
div
idu
als
Fre
qu
en
cy
Plegaderus caesus 652 X Hollow O 1 1 1 1 14 2
Gnathoncus nannetensis 672 Nest F 8 4 0 0 1 1
Gnathoncus buyssoni 674 Nest F 31 12 13 7 11 3
Gnathoncus buyssoni/ nannetensis
674/ 672 Nest F 5 3 2 2 0 0
Dendrophilus corticalis 677 Nest F 14 6 15 7 25 4
Paromalus flavicornis 680 Rotting O 5 1 3 2 1 1
Margarinotus striola 684 F 2 2 2 1 1 1
Acrotrichis insularis 784 1 1 0 0 0 0
Leiodes gyllenhali 836 0 0 1 1 0 0
Nemadus colonoides 888 X Nest O 16 10 29 10 20 10
Stenichnus godarti 948 Hollow O 0 0 0 0 2 2
Stenichnus bicolor 950 F 0 0 0 0 1 1
Euconnus pragensis 955 0 0 0 0 1 1
Scydmaenus hellwigi 965 Nest F 0 0 1 1 2 1
Gabrius splendidulus 994 F 6 2 0 0 2 1
Philonthus subuliformis 1030 127 17 13 6 17 6
Velleius dilatatus 1101 X Nest F 13 3 25 5 1 1
Quedius mesomelinus 1105 F 0 0 1 1 4 2
Quedius maurus 1106 Nest 1 1 0 0 0 0
Quedius cruentus 1107 F 0 0 0 0 3 3
Quedius brevicornis 1109 Nest O 144 16 17 11 14 3
Quedius scitus 1117 Nest F 2 2 1 1 0 0
Quedius xanthopus 1118 F 0 0 0 0 1 1
Othius melanocephalus 1174 0 0 0 0 1 1
Bibloporus bicolor 1330 O 0 0 1 1 1 1
Euplectus nanus 1338 Hollow F 0 0 0 0 17 5
Euplectus punctatus 1347 Rotting O 0 0 2 2 0 0
Euplectus karsteni 1349 Hollow F 1 1 12 4 127 20
Euplectus fauveli 1350 0 0 16 4 25 12
Hapalaraea melanocephala 1412 F 5 3 7 4 37 4
Hapalaraea nigra 1414 F 4 2 0 0 103 4
Hapalaraea floralis 1416 F 0 0 0 0 5 2
Hapalaraea ioptera 1421 O 0 0 0 0 2 1
Hapalaraea pygmaea 1424 X Nest O 5 3 2 2 0 0
1 Lundberg S (1995) Catalogus colepterorum. Naturhistoriska riksmuseet & Entomologiska föreningen i
Stockholm. 2 Gärdenfors U (ed) (2000) The 2000 red list of Swedish species. Artdatabanken SLU Sweden.
3 Ranius T & Jansson N (2002) A comparison of three methods to survey saproxylic beetles in hollow
oaks. Biodiversity and Conservation 11, 1759-1771.
23
Appendix B, Table 1 continued
Sp
ec
ies
Ca
t. N
o.
1
Sp
ec
ies o
f c
on
se
rva
tio
n
inte
res
t 2
Tre
ath
ca
teg
ory
Sw
ed
ish
Re
d l
ist
20
05
En
vir
on
men
t 3
Sa
pro
xy
lic
Fac
ult
ati
ve
=F
Sa
pro
xy
lic
Ob
lig
ate
=O
3
Ye
ar
1
Ye
ar
2
Ye
ar
3
No
. o
f in
div
idu
als
Fre
qu
en
cy
No
. o
f in
div
idu
als
Fre
qu
en
cy
No
. o
f in
div
idu
als
Fre
qu
en
cy
Sepedophilus testaceus 1635 F 13 7 0 0 0 0
Oxypoda soror 1744 1 1 121 15 0 0
Haploglossa gentilis 1781 X F 0 0 3 3 1 1
Haploglossa villosula 1782 F 106 13 0 0 38 6
Haploglossa marginalis 1785 F 4 2 1 1 0 0
Pentanota meuseli 1795 X NT O 1 1 0 0 0 0
Mocyta fungi 1920 0 0 0 0 0 0
Atheta crassicornis 1994 F 0 0 46 20 9 2
Atheta euryptera 1998 F 0 0 0 0 1 1
Atetha nigricornis 2001 F 554 28 13 2 192 20
Thamiaraea cinamomea 2054 O 5 2 1 1 0 0
Thamiaraea hospita 2055 X NT O 41 8 29 11 17 4
Prionocyphon serricornis 2197 X O 0 0 7 7 0 0
Trox scaber 2203 Nest O 28 10 1 1 48 10
Liocola marmorata 2291 X Hollow O 3 2 1 1 5 4
Potosia cuprea 2292 0 0 1 1 0 0
Osmoderma eremita 2293 X NT O 0 0 2 1 0 0
Selatosomus aeneus 2430 2 2 5 5 1 1
Ampedus pomonae 2438 O 0 0 0 0 0 0
Ampedus nigroflavus 2441 X NT Rotting O 2 2 1 1 4 4
Ampedus pomorum 2442 Rotting O 0 0 0 0 5 2
Ampedus hjorti 2443 X Hollow O 1 1 0 0 2 2
Ampedus balteuatus 2447 Rotting O 0 0 0 0 5 3
Ampedus tristis 2451 O 0 0 0 0 1 1
Ampedus nigrinus 2453 O 0 0 1 1 2 2
Melanotus castanipes 2459 Rotting O 0 0 0 0 1 1
Aulonothroscus brevicollis 2489 F 0 0 1 1 1 1
Trixagus dermestoides 2490 0 0 4 2 1 1
Trixagus carinifrons 2491 0 0 1 1 3 3
Dermestes lardarius 2562 Nest F 2 1 4 4 2 1
Globicornis emarginata 2567 F 3 1 1 1 0 0
Megatoma undata 2579 Rotting F 3 3 3 1 4 4
Ctesias serra 2581 Nest F 0 0 2 2 2 2
Anthrenus scrophulariae 2583 Nest F 1 1 1 1 2 1
Anthrenus museorum 2585 Nest F 1 1 1 1 4 3
Lyctus linearis 2592 X VU Dry O 0 0 0 0 2 1
Tipnus unicolor 2612 F 0 0 347 16 0 0
Ptinus rufipes 2617 Rotting O 0 0 13 6 1 1
Ptinus fur 2619 Hollow F 117 7 15 5 109 16
1 Lundberg S (1995) Catalogus colepterorum. Naturhistoriska riksmuseet & Entomologiska föreningen i
Stockholm. 2 Gärdenfors U (ed) (2000) The 2000 red list of Swedish species. Artdatabanken SLU Sweden.
3 Ranius T & Jansson N (2002) A comparison of three methods to survey saproxylic beetles in hollow
oaks. Biodiversity and Conservation 11, 1759-1771.
24
Appendix B, Table 1 continued
Sp
ec
ies
Ca
t. N
o.
1
Sp
ec
ies o
f c
on
se
rva
tio
n
inte
res
t 2
Tre
ath
ca
teg
ory
Sw
ed
ish
Re
d l
ist
20
05
En
vir
on
men
t 3
Sa
pro
xy
lic
Fac
ult
ati
ve
=F
Sa
pro
xy
lic
Ob
lig
ate
=O
3
Ye
ar
1
Ye
ar
2
Ye
ar
3
No
. o
f in
div
idu
als
Fre
qu
en
cy
No
. o
f in
div
idu
als
Fre
qu
en
cy
No
. o
f in
div
idu
als
Fre
qu
en
cy
Ptinus subpilosus 2622 Rotting O 7 3 0 0 20 12
Xestobium rufovillosum 2628 Dry O 0 0 0 0 2 2
Gastrallus immarginatus 2640 X dry O 0 0 1 1 2 1
Lymexylon navale 2675 X NT Dry O 7 4 1 1 3 2
Ostoma ferruginea 2679 O 0 0 0 0 0 0
Grynocharis oblonga 2682 X Rotting O 1 1 1 1 0 0
Hypebaeus flavipes 2716 X VU Rotting O 0 0 0 0 5 3
Glischrochilus hortensis 2837 O 5 2 0 0 0 0
Rhizophagus bipustulatus 2852 O 8 2 1 1 0 0
Rhizophagus cribratus 2856 Rotting O 0 0 0 0 1 1
Monotoma angusticollis 2859 0 0 2 1 0 0
Cryptophagus acutangulus 2907 F 0 0 10 2 1 1
Cryptophagus badius 2912 Rotting O 1 1 21 1 1 1
Cryptophagus micaceus 2922 X Nest O 1 1 0 0 62 4
Cryptophagus confusus 2926 X Hollow F 0 0 63 13 11 3
Cryptophagus dentatus 2928 Fungi F 1 1 0 0 5 3
Cryptophagus scanicus 2933 Rotting F 9 7 0 0 27 12
Cryptophagus pallidus 2934 X F 0 0 2 1 2 1
Cryptophagus scutellatus 2937 F 0 0 4 3 1 1
Atomaria morio 2953 Nest F 2 2 9 6 2 1
Dacne bipustulata 3015 O 0 0 0 0 2 1
Cerylon histeroides 3037 Rotting O 24 10 2 1 1 1
Cerylon ferrugineum 3038 Rotting O 6 3 2 2 0 0
Latridius hirtus 3134 O 4 3 1 1 0 0
Latridius consimilis 3135 F 6 4 0 0 1 1
Latridius minutus 3137 F 0 0 2 2 2 1
Latridius nidicola 3139 F 1 1 0 0 2 1
Enicmus rugosus 3146 O 6 6 1 1 1 1
Dienerella elongata 3150 F 5 2 1 1 0 0
Cartodere separanda 3151 5 3 44 12 0 0
Cartodere constricta 3168 F 1 1 2 2 0 0
Corticaria serrata 3179 F 58 11 4 1 11 5
Corticaria longicollis 3188 F 5 3 2 1 0 0
Corticarina elongata 3193 F 0 0 0 0 0 0
Mycetophagus piceus 3261 X Rotting O 0 0 0 0 0 0
Mycetophagus 4-guttatus 3264 X VU Fungi F 2 1 0 0 18 1
Diaperis boleti 3343 Fungi O 1 1 2 2 1 1
Alphitobius diaperinus 3370 F 1 1 2 1 0 0
Tenebrio opacus 3383 X VU Hollow O 4 1 20 11 1 1
1 Lundberg S (1995) Catalogus colepterorum. Naturhistoriska riksmuseet & Entomologiska föreningen i
Stockholm. 2 Gärdenfors U (ed) (2000) The 2000 red list of Swedish species. Artdatabanken SLU Sweden.
3 Ranius T & Jansson N (2002) A comparison of three methods to survey saproxylic beetles in hollow
oaks. Biodiversity and Conservation 11, 1759-1771.
25
Appendix B, Table 1 continued
Sp
ec
ies
Ca
t. N
o.
1
Sp
ec
ies o
f c
on
se
rva
tio
n
inte
res
t 2
Tre
ath
ca
teg
ory
Sw
ed
ish
Re
d l
ist
20
05
En
vir
on
men
t 3
Sa
pro
xy
lic
Fac
ult
ati
ve
=F
Sa
pro
xy
lic
Ob
lig
ate
=O
3
Ye
ar
1
Ye
ar
2
Ye
ar
3
No
. o
f in
div
idu
als
Fre
qu
en
cy
No
. o
f in
div
idu
als
Fre
qu
en
cy
No
. o
f in
div
idu
als
Fre
qu
en
cy
Tenebrio molitor 3385 Hollow F 6 1 0 0 0 0
Prionychus ater 3400 Hollow O 9 5 0 0 23 8
Pseudocistela ceramboides 3403 Hollow O 3 1 13 4 0 0
Mycetochara humeralis 3408 X NT Hollow O 0 0 0 0 2 2
Mycetochara linearis 3410 Rotting O 8 3 0 0 6 3
Scraptia fuscula 3414 X Nest O 0 0 1 1 42 10
Orchesia undulata 3463 Fungi O 2 2 11 1 0 0
Xyleborinus saxesenii 4516 X NT O 0 0 1703 14 0 0
Lasius brunneus 5002 1 1 129 2 7 1
Lasius niger 5003 1024 10 778 5 1087 18
Lasius fuliginosus 5004 122 2 3 2 342 2
Formica rufa grp 5005 750 6 1 1 346 5
Camponotus sp 5006 58 9 2 2 0 0
Allochernes wideri 0 0 0 0 0 0
Anthrenochernes stellae X 3 2 9 4 0 0
Apocheiridium ferum DD 0 0 3 3 1 1
Brachypalpus laphriformis NT 5 5 32 16 0 0
Chernes cimicoides NT 2 2 9 7 1 1
Ctenophora pectinicornis 32 14 11 6 48 14
Dictendia bimaculata 6 6 45 17 13 7
Dinocheirus panzeri 1 1 60 20 5 3
Geting mindre 22 15 2 2 22 11
Vespa crabro 23 11 3 2 5 4
Mytharopa florea 7 6 0 0 1 1
Pocota personata X 2 2 0 0 0 0
1 Lundberg S (1995) Catalogus colepterorum. Naturhistoriska riksmuseet & Entomologiska föreningen i
Stockholm. 2 Gärdenfors U (ed) (2000) The 2000 red list of Swedish species. Artdatabanken SLU Sweden.
3 Ranius T & Jansson N (2002) A comparison of three methods to survey saproxylic beetles in hollow
oaks. Biodiversity and Conservation 11, 1759-1771.