Evaluation of artificial habitats for saproxylic oak …liu.diva-portal.org/smash/get/diva2:18114/FULLTEXT01.pdfbeetles (Palm 1959). The definition of saproxylic species is species

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

7 References

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17

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saproxylic beetles in hollow oaks. Biodiversity and Conservation 11, 1759-

1771.

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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 - ***


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 + *


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


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 - ***


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


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 - ***


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


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


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


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





24


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





25


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



