Inventory report on insects
depending on logs of
Pinus sylvestris
S:t Anna, Östergötland, Sweden
Final report
Photo Karl-Olof Bergman
By:
Sara Eklund
Emma Larsson
University of Linköping
20040301
Sarek inventering
Anitys – Inventering av skog & mark
Contents
1 Abstract...........................................................................................................................1
2 Introduction.....................................................................................................................1
3 Material and methods..................................................................................................... 3
3.1 Study area................................................................................................................3
3.2 Selection of research sites.......................................................................................3
3.3 Inventory of beetles.................................................................................................3
3.4 Measurements of environmental variables.............................................................4
3.4.1 Tree scale.........................................................................................................4
3.4.2 Landscape scale...............................................................................................4
3.5 Statistics..................................................................................................................5
3.5.1 Tree scale.........................................................................................................5
4 Site descriptions..............................................................................................................6
4.1 Röviken ..................................................................................................................6
4.2 Sanningsholmen .....................................................................................................6
4.3 Ugglö ......................................................................................................................6
4.4 Yxnö .......................................................................................................................7
4.5 Långholmen ...........................................................................................................7
4.6 Grytsholm ...............................................................................................................7
4.7 Hästö ......................................................................................................................7
4.8 Törnsholm ..............................................................................................................7
4.9 Fågelö .....................................................................................................................7
4.10 Gröskär .................................................................................................................8
4.11 Yxnerum ..............................................................................................................8
5 Results and discussion.................................................................................................... 8
5.1 Tree scale................................................................................................................9
5.2 Landscape scale.....................................................................................................12
6 Conclusion.................................................................................................................... 16
7 Acknowledgements.......................................................................................................16
8 References.....................................................................................................................17
Appendix.........................................................................................................................20
1 Abstract
Variables affecting communities of saproxylic beetle species, depending on logs of
Pinus sylvestris L. were investigated in this study. The study took place on ten islands in
the archipelago of S:t Anna, Östergötland, south-eastern Sweden. One site, situated on
the mainland of Östergötland, was used as a reference. The aim of the study was to
investigate factors affecting the species community and species richness of saproxylic
beetles dependent on Pinus sylvestris. Estimations of factors were made on two
different scales; tree and landscape. On tree scale the structure of the beetle communities
differed to some extent and sun exposure was found to be the most important factor.
Red-listed species showed a higher positive correlation with log diameter than
saproxylic species in general. On landscape scale, both the number of species and the
presence of red-listed species increased significantly with increasing proportion of
coniferous forest in the surroundings. The conclusion is that in conserving saproxylic
beetle species it is vital to maintain large areas of coniferous forest with a high
variability in structure.
2 Introduction
Sweden is to 55% covered with forest and 85% of this is dominated by the coniferous
trees Pinus sylvestris L. and Picea abies L.. Forestry has for the past centuries been
important to the Swedish economy (Engelmark & Hytteborn 1999). The production of
wood and paper has left little of the forest areas unaffected (Ehnström & Waldén 1986)
and even the forest in the archipelago has been affected to a great extent. The
archipelago is a unique type of nature, which is mainly found along the Scandinavian
coast. Therefore Sweden has a responsibility to protect these areas.
Due to the large negative affects of forestry on biodiversity, a valuation system
named woodland key habitats has been created to define areas with natural forest
structures and presence of red-listed species (Engelmark & Hytteborn 1999). Today only
0.6% of the forest in Sweden is considered woodland key habitats according to the
Swedish National Board of Forestry (2003). Another result of the intensive forest
management is the decreased amount of coarse woody debris (CWD) (Jonsell et al.
1998, Similä et al. 2003). This decrease has been documented having considerable
effects on species diversity in the forests (Jonsell et al 1998, Schiegg 2000b). Siitonen
(2001) estimates that 20-25% of all forest dwelling species depend of dead wood.
Beetles, which is one of the largest groups of saproxylic insects in Sweden (Kaila et al.
1997), will to a great extent be affected by a decreasing amount of CWD.
According to Jonsell et al. (1998) pine forest is an important biotope because of the
large number of saproxylic species that depend on Pinus sylvestris. Today's reduced
amount of sites with natural pine forests has led to a fragmented landscape for a number
of beetles. Increased distances between suitable sites and a reduced area size results in
smaller populations that are more sensitive to environmental changes. The increased
isolation decreases the probability of recolonization (Thomas et al 1992, Økland et al.
1996). Fragmentation of the forest in the archipelago will have an even greater affect on
the species community, due to the already fragmented landscape that the archipelago
exhibit. Apart from the fragmentation of pine forest at landscape scale, fragmentation
can also be observed at site scale as increasing distances between single dead trees
within the forest (Schiegg 2000a, 2000b). Studies have shown that the connectivity of
1
CWD is more important than the quantity of the substrate (Rukke 2000, Schiegg 2000b).
The quality of the log has also been documented to affect the beetle community (Wikars
2003).
The main aim of this study was to investigate the effects of surrounding landscape
on species richness of beetles. The study was based on factors affecting beetle species
depending on logs of Pinus sylvestris. Another aim was to investigate how the quality of
the log, e.g. sun exposure, diameter and decaying stage, affect the beetle community.
The inventory was made in collaboration with the University of Linköping and the Life
project – Coastal Woodlands.
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4
10
5
3
1
8
2
7
2
9
3 Material and methods
3.1 Study area
The study was conducted on islands in the archipelago of S:t Anna, 40 km south-east of
Söderköping, Sweden (58°20N, 16°55E) (Figure 1). The archipelago consists of islands
of different sizes, some covered by sparse vegetation others only consisting of barren
rocks. One area, situated on the mainland of Östergötland, was used as a reference. The
forest in all areas is dominated by Pinus sylvestris.
3.2 Selection of research sites
Sites in the archipelago dominated by Pinus sylvestris were identified using GIS
(ArcView 3.2) with data from the Natura 2000 and the woodland key habitat
inventories. According to the Swedish National Board of Forestry (2003), a woodland
key habitat is an area within the forest having qualities that support rare and endangered
species with highly specific demands of habitat. All sites selected in the area had Pinus
sylvestris as first or second most dominating tree species. From these sites, ten research
sites were chosen with similar conditions regarding log frequency and size. The research
sites had an area range between 14000-39000 m2 and an estimated proportion of
coniferous forest in the surrounding area (radius 1000 m) between 1-56%. Sites were
also chosen to have a different degree of connectivity, with respect to woodland key
habitats and Natura 2000 sites. The connectivity (Si) was calculated using a formula
created by Hanski (1999).
Si = ∑ exp (-αdij) Aj,
The formula includes the areas (Aj) of surrounding sites and the distances (dij) to these.
The α-value is a constant and α = 0.0027 was used in the calculations. This value means
that 6% of the population will reach a distance of 1000 m. The value was chosen in
consideration to saproxylic beetle's limited ability to spread (Warren & Key 1991) and a
study made on Galerucella calmariensis (L.) by Grevstad & Herzig (1997). The
reference site on the mainland was chosen to be an area having equally good qualities as
the sites in the archipelago. This was made to facilitate the comparison between the
archipelago and the mainland.
3.3 Inventory of beetles
Eight window traps were placed on logs of Pinus sylvestris on each research site. The
traps consisted of a transparent plastic pane (35*60 cm) with an aluminium tray
(13*33*17 cm) hanging underneath. The tray contained a 1.5-litre solution of 60%
glycol, 39% water, detergent and Bitrex. The detergent eliminated surface tension and
Bitrex made the solution distasteful to mammals. Attached to the plastic pane were two
test tubes containing turpentine and ethanol (95%), which were used to attract beetles
associated with newly dead wood (Schroeder 1988). The inventory period stretched
from the 23rd of April to the 5th of September and the traps were emptied four times
during this period, once a month. All beetle species were identified to species level and
the naming followed Catalogus Coleopterorum Sueciae (Lundberg 1995).
3
Figure 2 Window trap on the research site Röviken in the archipelago of S:t Anna,
Östergötland. Photo Emma Larsson.
3.4 Measurements of environmental variables
Environmental variables were measured on the research sites on two spatial scales; tree
and landscape scale (Table 1). These variables were thought to affect the beetle fauna
depending on logs of Pinus sylvestris.
3.4.1 Tree scale
Estimations on tree scale were made on the appearance of the log, as log diameter (in
meters) and quality. The decaying stage of the log was an estimation of log quality and
was divided into four categories; parts of the bark left, silver log, external wood decayed
and external and internal wood decayed. Additional estimations were made on the
spatial direction of the log (north, north-east, east, south-east), which was then
transformed to a continuous variable using the cosines function. The ground type was
noted as rock or soil and estimations were made on the percentage of ground contact.
The logs exposure to the sun was estimated in two ways; how large part of the day the
log was exposed to the sun and how large part of the log that was not exposed to the sun
at all during the day. These two estimations were then used to create an index for the
total sun exposure of the log. The index, which was later used in the statistical analysis,
was created through multiplying the two percentage values.
3.4.2 Landscape scale
The quantity of logs and snags were estimated in all research sites. The area of the
2
research sites were estimated in m and a general description was made of each site.
Connectivity was a measurement of isolation on landscape scale, together with
estimations of the proportion of coniferous forest and the proportion of land surrounding
the research sites.
4
Table 1 The environmental variables estimated on tree and landscape scale and used in the
statistical analysis.
Environmental variables
Tree scale
Landscape scale
Sun exposure index
Total number of species
Ground contact
Number of red-listed species
Ground type
Number of logs
Direction of the log
Density of logs
Decay stage of the log
Number of snags
Log diameter
Density of snags
Area of the research site
Connectivity
Proportion of coniferous forest in the
surrounding area (radius 1000 m)
Proportion of land in the surrounding area
(radius 1000 m)
3.5 Statistics
The statistical analysis were made on a selection of species caught in the study. The
selected species were those depending on coniferous trees and the categorisation was
made in consultation with the entomologists Gunnar Sjödin1 and Nicklas Jansson2 and
by using Palm (1959) and Ehnström (2002). On tree scale individuals from the same
trap were summarized and used as a sample. Six of the traps in the study failed to
produce any results, therefore 74 samples were used in the statistical analysis on the data
from the archipelago. The samples on landscape scales were a summary of all traps at
each research site.
3.5.1 Tree scale
The statistical program Canoco 4.5 was used to make multivariate statistical analysis
(ter Braak & Šmilauer 2002, Lepš & Šmilauer 2003). The data on tree scale were first
tested with a detrended correspondence analysis (DCA) to calculate the length of the
gradients. The analysis showed that the length of the gradient was 2.163, which is less
than 4. This therefore allowed for further linear analysis (Lepš & Šmilauer 2003). A
partial redundancy analysis (partial RDA) was made to analyse the relationship between
species and environmental variables (Table 1). The ten research sites acted as ten
categorical covariables in the analysis and a Monte Carlo test was used to test for
statistical significance of all canonical axes, using 9999 permutations. The relationship
between sun exposure and both the total number of species and the number of red-listed
species was also tested with linear regression.
1 Gunnar Sjödin, entomologist
2 Nicklas Jansson, IFM, University of Linköping, Sweden
5
Figure 3 Prionus coriarius (L.) found in the archipelago of S:t Anna, Östergötland, and
categorized as near threatned on the Swedish red-list. Photo Karl-Olof Bergman.
4 Site descriptions
4.1 Röviken
The site was an elongated part of a mountain in an east-west direction, with a steep
rocky slope facing the Baltic See to the south. The site consisted mainly of barren rock
and had a small moist area in the centre covered by Vaccinium myrtillus L.. There were
about 70 logs, exclusively silver logs, of Pinus sylvestris at the site and about 30
standing dead trees.
(Area size 2.9 ha)
4.2 Sanningsholmen
The site consisted to some extent of barren rocks and scattered over the area were Picea
abies and the deciduous trees Quercus robur L. and Populus tremula L.. The northern
part of the area was rather moist and the ground vegetation to the east was dominated by
Vaccinium myrtillus. There were about 20 logs, mainly silver logs, of Pinus sylvestris at
the site and about 10 standing dead trees.
(Area size 1.6 ha)
4.3 Ugglö
This site was a mountain with a steep rocky side facing south-west and a main part
consisting of barren rocks. The centre of the site contained deciduous trees like Quercus
robur and Betula pendula L. and the ground vegetation dominated by Vaccinium
myrtillus and Vaccinium vitis-idaea L. The site was surrounded by several Quercus
robur and some trees were fairly large. There were about 40 logs, mainly silver logs, of
Pinus sylvestris and about 20 standing dead trees.
(Area size 1.4 ha)
6
4.4 Yxnö
Yxnö is a research site that mainly consisted of barren rocks on a west side of a
mountain. A moist elongated area, with Picea abies and Quercus robur, divided the
slope in two. Juniperus communis L. was found in the lower parts of the slope, while the
upper part mainly consisted of Pinus sylvestris. There were about 60 logs, mainly silver
logs, of Pinus sylvestris at the site and about five standing dead trees.
(Area size 2.6 ha)
4.5 Långholmen
An elongated mountain ridge in an east-west direction formed this research site. The
ground was to a great extent covered by Leucobryum glaucum L. and Pinus sylvestris
was the only tree species present. There were about 50 logs, mainly logs with the
external wood decayed, and about 20 standing dead trees of Pinus sylvestris at the site.
(Area size 2.4 ha)
4.6 Grytsholm
The west part of the island consisted of barren rocks while the east part was more closed
with a vegetation of grass, bushes and deciduous trees. The dominating tree species was
Pinus sylvestris, but the site also contained several large Tilia cordata Mill. and some
Quercus robur and Sorbus aucuparia L.. There were about 25 logs, mainly silver logs,
and about five standing dead trees of Pinus sylvestris at the site.
(Area size 2.1 ha)
4.7 Hästö
This site had a varying vegetation with open parts consisting of rocks covered by
Cladonia spp. The open parts were separated by closed moist areas dominated by Betula
pendula and Sorbus aucuparia. There were about 20 logs, mainly silver logs, of Pinus
sylvestris at the site and about 5 standing dead trees.
(Area size 3.9 ha)
4.8 Törnsholm
The site was an elongated mountain in a north-south direction with a steep rocky side
facing east. Moist parts with Vaccinium myrtillus were found between areas of barren
rocks. The dominating tree vegetation besides Pinus sylvestris were Quercus robur and
Juniperus communis. There were about 35 logs, mainly silver logs, of Pinus sylvestris at
the site and about 20 standing dead trees.
(Area size 2.6 ha)
4.9 Fågelö
The elongated mountain ridge had a steep rocky side facing west. The site consisted
mainly of barren rocks but had a moist area, dominated by Picea abies and Betula
pendula, that separates the site in a northern and southern part. There were about 40
logs, mainly silver logs and logs with the external wood decayed, of Pinus sylvestris at
the site and about 20 standing dead trees.
(Area size 2.6 ha)
7
4.10 Gröskär
The island has a closed centre part with a large number of Juniperus communis and the
island edges consisted of barren rocks. A moist area dominated by Pteridium aquilinum
L. was found in the centre of the site. There were about 15 logs, mainly logs with bark
left and logs with external wood decayed, of Pinus sylvestris found at the site and about
five standing dead trees.
(Area size 1.5 ha)
4.11 Yxnerum
This site was situated 40 kilometres south east of Linköping and surrounded by several
woodland key habitats dominated by coniferous forest. The research site was a mountain
with steep rocky sides facing south and east. Cladonia spp. covered the rocks and the
tree vegetation was exclusively made up by Pinus sylvestris. There were about 30 logs,
mainly silver logs, of Pinus sylvestris at the site and about 10 standing dead trees.
(Area size 1.8 ha)
5 Results and discussion
In total, 14500 individuals of 563 beetle species were caught in our study. These species
counts for 12% of the total Swedish coleopteran fauna (Wastenson 1996). Of the total
number of species were 210 (37%) species saproxylic and associated with coniferous
trees (Palm 1959, Ehnström 2002, personal communication Nicklas Jansson and Gunnar
Sjödin). Of these, 15 species (210 individuals) are red-listed in Sweden. Ten of the redlisted species were categorized as near-threatened and five as vulnerable according to
the Swedish red-list (Artdatabanken 2003).
Only few individuals of the red-listed species were found, except Stagetus borealis
(Israelsson) which was found in almost all research sites and with as much as 28
individuals on Yxnö. Single individuals of Ebaeus lapplandicus (Evers), Ptinus
sexpunctatus (Panzer), Stenagostus rufus (DeGeer), Grynocharis oblonga (L.) and
Ipidia binotata (Reitter), categorised as vulnerable according to the Swedish red-list
(Artdatabanken 2003), were found in the study area. Prionychus melanarius (Germar),
categorised as an endangered species, has been documented to be associated with
deciduous trees as Betula pendula, Quercus robur and Fagus sylvatica L. Despite the
lack of these tree species in several of the study sites Prionychus melanarius were found
in seven of the 10 research sites in the archipelago of St Anna. The largest number of
species and red-listed species in the archipelago was found at Ugglö and the lowest
number at Gröskär and Fågelö (Table 2).
Several of the sites in the archipelago were found having the same high quality as
Yxnerum, which was a site surrounded by a large proportion of coniferous forest and
woodland key habitats. Four sites in the archipelago had an average number of red-listed
species that was as large as the number of species found at Yxnerum.
8
Table 2 The total number of species, the number of individuals and the number of red-listed
species on each research site. The numbers in parenthesis show the average number per trap.
Sites
Number of species
Number of
Number of red-listed
individuals
species
Total
Average
Total
Average
Total
Average
1 Röviken
108
(13.5)
1019
(127.4)
8
(1.00)
2 Sanningsholmen
86
(17.2)
590
(118.0)
5
(1.00)
3 Ugglö
114
(14.3)
1361
(170.1)
10
(1.25)
4 Yxnö
94
(13.4)
870
(124.3)
8
(1.14)
5 Långholmen
104
(13.0)
1212
(151.5)
5
(0.63)
6 Grytsholm
94
(11.8)
821
(102.6)
3
(0.38)
7 Hästö
93
(13.3)
786
(112.3)
4
(0.57)
8 Törnsholm
73
(9.1)
526
(65.8)
3
(0.38)
9 Fågelö
70
(8.8)
508
(63.5)
2
(0.25)
10 Gröskär
70
(10.0)
562
(80.3)
1
(0.14)
125
(15.6)
2334
(291.8)
8
(1.00)
Archipelago
Mainland
11 Yxnerum
5.1 Tree scale
The statistics showed that all environmental variables together affected the beetle
community significantly (partial RDA, Monte Carlo test: Sum of eigenvalues = 0.123,
F = 1.986, p = 0.0021). Sun exposure was the environmental variable that best could
explain the beetle community of the logs. The results showed that there was one group
of species positively correlated with sun exposure and another large group with a strong
negative correlation. Of the species documented to mainly depend on dead wood of
Pinus sylvestris (Ehnström 2002), 50% of the species in our study were found to be
negatively correlated with sun exposure. This result contradicts the general theory that
saproxylic species, dependent on Pinus sylvestris, are associated with high sun exposure
(Cederberg et al. 1997, Ehnström 2002, Wikars 2003). These results indicate the
importance of a range of logs with different grade of sun exposure in the forest when
preserving a rich beetle fauna. Another result regarding the species having a strong
correlation with shaded conditions is that several of these species are associated with
fungus (personal communication Gunnar Sjödin and Nicklas Jansson). Fungus often
thrive in the shade and is favoured by high humidity.
Further analysis, using linear regression, showed that sun exposure did not affect
the number of species per trap (F1,72 = 1.111, p = 0.295) nor the number of red-listed
species (F1,72 = 0.239, p = 0.626). Despite these results sun exposure were found to affect
the structure of the beetle community. This result is in line with Sverdrup-Thygeson &
Ims (2002) study on saproxylic beetles on dead wood of Populus tremula.
9
Figure 4 Examples of species found
to be affected by sun exposure and
log diameter were
Anoplodera rubra,
Ebaeus lapplandicus (VU),
Ennearthron cornutum,
Hadreule elongatula,
Ptinus sexpunctatus (VU) and
Thanasimus femoralis.
Illustration Emma Larsson.
Figure 5 Examples of species found
to be affected by log diameter and
thrive in the shade were
Anaspis thoracica,
Arpidiphorus orbiculatus,
Athous haemorrhoidalis,
Athous subfuscus,
Cis alter,
Cis hispidus,
Enicmus rugosus,
Sphindus dubius and
Strophosoma capitatum.
Illustration Emma Larsson.
10
In the analysis of the red-listed species the amount of sun exposure were also found
to be important. Some species were associated with high sun exposure and others
thrived in shaded conditions. A large log diameter was another variable which had a
positive correlation with several of the red-listed beetle species. These results
correspond with a study on Tragosoma depsarium (L.) (Wikars 2003), which showed
that a large log diameter was important for the presence of the species. Stokland (2001)
found similar results on log diameter, but also found the presence of logs of different
decaying stages to be important. This imply that logs with large diameter are of
importance to maintain a high diversity of the beetle fauna. If only logs with a small
diameter are left in the forest, there will be a lack of substrate for species depending on
larger diameter. On the other hand if the forest contain logs with large diameters this
substrate will satisfy species with different preference in log diameter.
The beetle fauna in the log changes over time as a result of the decaying stage. We
found in the analysis that the decay of the log could to some extent explain the beetle
fauna present. Half of our research sites had less than two newly dead logs, i.e. logs with
bark left, but despite this low amount of newly dead wood species like Epuraea spp.,
Glischrochilus quadripunctatus (L.), Hylastes brunneus (Er.), Hylastes opacus (Er.),
Hylurgops palliatus (Gyll.), Rhizophagus depressus (F.), Thanasimus formicarius (L.)
and Tomicus piniperda (L.) were found. These species have been documented being
attracted by the scent of ethanol and α-pinene (Schroeder 1988, Schroeder & Lindelöw
1989, Byers et al. 2000), which is emitted from newly dead wood. These findings are
contradictory because of the limited amount of this substrate. The presence of the scent
traps baited with ethanol and turpentine could explain these results, which indicate that
these species have a rather good ability to spread. Species that depend on later stages of
succession are on the other hand generally thought to have a poor ability to spread.
Therefore a continuity of logs of different decaying stages is important.
Figure 6 Buprestis octoguttata (L.) found in the archipelago of S:t Anna and rated
as vulnerable on the Swedish red-list. Photo Karl-Olof Bergman.
11
5.2 Landscape scale
The most important factor on landscape scale affecting species richness of beetles was
the proportion of coniferous forest in the surrounding area. We found a positive
correlation between this factor and both the number of species (F1,8 = 10.901, p = 0.013)
and the number of red-listed species (F1,8 = 77.399, p = 0.00005) (Figure 8). These
results are similar to those found by Burke & Goulet (1998) on the effect of forest
fragmentation on beetle biomass. This also imply that increasing isolation has negative
effects on species communities (Økland et al. 1996, Rukke 2000). Increasing distance to
the mainland was another factor having a negative correlation with the number of
species. Similar results, on the importance of distance, were found in a study on carabid
beetles by Kotze & Niemelä (2002). In their study a greater number of species was
found on the mainland of Åland compared to the smaller islands in the archipelago. The
results of both these studies are in accordance with the island biogeography theory
(Akçakaya et al. 1999). However, as the proportion of coniferous forest around the sites
decreases with increasing distance to the mainland, it is not possible to determine which
of these factors that are responsible for the pattern of species richness.
Figure 7 The archipelago of S:t Anna in the summer of 2003. Photo Karl-Olof Bergman.
12
Total number of species
20
2
15
7
5
4
6
10
10
9
8
5
0
0.0
Number of red-listed species
3
1
0.2
0.4
0.6
1.5
3
1 2
1.0
4
5
7
0.5
8
10
0.0
0.0
6
9
0.2
0.4
0.6
Proportion of coniferous forest
Figure 8 The relationship between the proportion of coniferous forest in the
surrounding area and the mean number of saproxylic beetles species per site,
depending on logs of Pinus sylvestris in the archipelago of S:t Anna.
13
Connectivity, a second estimation on isolation, based only on areas classified as
being of high quality according to red-listed species, did however not affect the number
of species (F1,8 = 0.137, p = 0.722) nor the number of red-listed species (F1,8 = 0.0001,
p = 0.992) (Figure 9). This may be a result of the surrounding areas, not included in the
connectivity estimation, to generally be of high quality and therefore have a great impact
on the beetles.
20
Total number of species
2
15
1
7
3
6
10
10
4
5
9
8
5
0
2.0
3.0
4.0
5.0
Red-listed species
2.0
1.5
1
7
3
4
2
1.0
5
6
0.5
10
0.0
2.0
3.0
Connectivity
4.0
8
9
5.0
Figure 9 The relationship between connectivity on the number of saproxylic beetle
species, depending on logs of Pinus sylvestris in the archipelago of S:t Anna.
14
The correlation matrix also show the importance of a high proportion of coniferous
forest in the surroundings on the presence of red-listed species (Table 3). Log density
affected the number of red-listed species, but could in our study not be differentiated
from the effects of the proportion of coniferous forest. Schiegg (2000a, 2000b) and
Stokland (2001) on the other hand found that the number of saproxylic species was
positively correlated with a high amount of CWD in the forest. The substrate is without
doubt crucial for these species, as saproxylic species are by definition associated with
dead wood (Speight 1989).
# spec
Prop. land
Prop. forest
Area
Con
D snags
# snags
D logs
# logs
Red-list
# spec
Table 3 The correlation matrix show the relationship between environmental variables. Numbers
in bold show relationships with significant correlation (p<0.05). The abbreviations in the matrix
are;1) Number of species, 2) Number of red-listed species, 3) Number of logs, 4) Density of
logs, 5) Number of snags, 6) Density of snags, 7) Connectivity, 8) Site area, 9) Proportion of
coniferous forest (radius 1000 m) and 10) Proportion of land (radius 1000 m). The correlation
matrix has not been compensated for multiple tests.
1
Red-list 0.796
1
# logs
0.086
0.486
1
D logs
0.267
0.699
0.826
1
# snags 0.107
0.354
0.633
0.644
1
D
snags
0.303
0.524
0.336
0.681
0.834
1
Con
-0.161
-0.021
-0.016
-0.004
-0.034
-0.022 1
Area
-0.168
-0.136
0.287
-0.207
-0.021
-0.440 0.011
1
Prop.
forest
0.690
0.936
0.528
0.745
0.285
0.445
0.134
-0.264
1
Prop.
land
0.575
0.887
0.543
0.707
0.157
0.292
0.155
-0.195
0.964
15
1
6 Conclusion
In conserving saproxylic species, associated with Pinus sylvestris, several factors should
be taken under consideration. The most important factor was a large proportion of
coniferous forest surrounding the site. This imply that conservation measures will have a
greater effect preventing forest fragmentation rather than protecting single sites of good
quality. Another factor of importance was the amount of CWD in the forest. The logs
should have a large diameter and be located in different degrees of sun exposure. Both
wetter shaded parts as well as dryer rocky parts are necessary in the forest to keep a high
species diversity.
7 Acknowledgements
First of all we want to thank our supervisor Karl-Olof Bergman at the University of
Linköping for all his help and support. We are grateful to Per Milberg at the University
of Linköping for helping with the statistical analysis. For transportation in the
archipelago we are much obliged to Mats Södling, without him we would have been
stranded. We are very thankful to Gunnar Sjödin, who identified all 520 beetle species,
and Nicklas Jansson at the University of Linköping for help with the classification. We
also want to thank our contacts Hans Liman at Sweden´s County Administration
Östergötland and Bo Thor at the Swedish National Board of Forestry. At last we want to
thank the Life-project Coastal woodlands for financial support.
Figure 10 Sara Eklund and Emma Larsson in the archipelago of S:t Anna, Östergötland. Photo
Karl-Olof Bergman.
16
8 References
Akçakaya HR, Burgman MA & Ginzburg LR (1999) Applied population ecology. 2nd ed.
Sinauer Associates Inc. Publishers, Sunderland, Massachusetts.
Artdatabanken, SLU, (2003) Rödlistade arter i Sverige 2000. http://wwwumea.slu.se/MiljoData/webrod/SOKNING.cfm (2nd January 2004)
Burke D & Goulet H (1998) Landscape and area effects on beetle assemblages in
Ontario. Ecography 21, 472-479.
Byers JA, Zhang QH & Birgersson G (2000) Strategies of a bark beetle, Pityogenes
bidentatus, in an olfactory landscape. Naturwissenschaften 87, 503-507.
Cederberg B, Ehnström B, Gärdenfors U, Hallingbäck T, Ingelög T & Tjernberg M
(1997) De trädbärande impedimentens betydelse för rödlistade arter. Artdatabanken,
SLU, Uppsala.
Ehnström B (2002) Insektsgnag i bark och ved. Artdatabanken, Uppsala.
Ehnström B & Waldén HW (1986) Faunavård i skogsbruket. D2, Den lägre faunan.
Skogsstyrelsen, Jönköping.
Engelmark O & Hytterborn H (1999) Coniferous forest. pp 55-74 in: Rydin H (ed.)
Swedish plant geography. Svenska växtgeografiska sällskapet, Uppsala.
Grevstad FS & Herzig L (1997) Quantifying the effects of distance and conspecifics on
colonization: experiments and models using the loosestrife leaf beetle, Galerucella
calmariensis. Oecologia 110, 60-68.
Hanski I (1999) Habitat connectivity, habitat continuity, and metapopulations in
dynamic landscapes. Oikos 87, 209-219.
Jonsell M, Weslien J & Ehnström B (1998) Substrate requirements of red-listed
saproxylic invertebrates in Sweden. Biodiversity and Conservation 7, 749-764.
Kaila L, Martikainen P & Punttila P (1997) Dead trees left in clear-cuts benefit saroxylic
Coleoptera adapted to natural disturbances in boreal forest. Biodiversity and
Consevation 6, 1-18.
Kotze DJ & Niemelä J (2002) Year-to-year variation in carabid beetle (Coleoptera,
Carabidae) assemblages on the Åland Islands, south-west Finland. Journal of
Biogeography 29, 375-386.
Lepš J & Šmilauer P (2003) Multivariate Analysis of Ecological data using CANOCO.
University Press, Cambridge.
17
Lundberg S (1995) Catalogus Coleopterorum Sueciae. Entomologiska föreningen,
Naturhistoriska riksmuseet, Stockholm.
Palm T (1959) Die Holz- und Rinden- Käfer der Süd- und Mittelschwedischen
Laubbäume. Lund.
Rukke BA (2000) Effects of habitat fragmentation: increased isolation and reduced
habitat size reduces the incidence of dead wood fungi beetles in a fragmented forest
landscape. Ecography 23, 492-502.
Schiegg K (2000a) Are there saproxylic beetle species characteristic of high dead wood
connectivity? Ecography 23, 579-587.
Schiegg K (2000b) Effects of dead wood volume and connectivity on saproxylic insect
species diversity. Ecoscience 7, 290-298.
Schroeder LM (1988) Attraction of the bark beetle Tomicus piniperda and some other
bark- and wood-living beetles to the host volatiles α-pinene and ethanol. Entomologia
Experimentalis et Applicata 46, 203-210.
Schroeder LM & Lindelöw Å (1989) Attraction of scolytids and associated beetles by
different absolute amounts and proportions of α-pinene and ethanol. Journal of
Chemical Ecology 15, 807-817.
Siitonen J (2001) Forest management, coarse woody debris and saproxylic organisms:
Fennoscandian boreal forests as an example. Ecological Bulletin 49, 11-41.
Similä M, Kouki J & Martikainen P (2003) Saproxylic beetles in managed and
seminatural Scots pine forests: quality of dead wood matters. Forest Ecology and
Management 174, 365-381.
Speight MCD (1989) Saproxylic invertebrates and their conservation. Council of
Europe, Strasbourg.
Stokland (2001) The coarse woody debris profile: an archive of recent forest history and
an important biodiversity indicator. Ecological Bulletins 49, 71-83.
Sverdrup-Thygeson A & Ims RA (2002) The effects of forest clearcutting in Norway on
the community of saproxylic beetles on aspen. Biological Conservation 106, 347-357.
Swedish National Board of Forestry (2003). http://www.svo.se (29th December 2003)
ter Braak CJF & Šmilauer P (2002) CANOCO 4.5. Biometris, Wageningen, the
Netherlands.
Thomas CD, Thomas JA & Warren MS (1992) Distributions of occupied and vacant
butterfly habitats in fragmented landscapes. Oecologia 92, 563-567.
18
Warren MS & Key RS (1991) Woodlands: past, present and potentials for insects. pp
155-211. In Collins N M & Thomas J A (ed.). The Conservation of Insects and their
Habitats, 15th symposium of the Royal Entomological Society of London. Academic
press, London.
Wastenson L (ed.) (1996) Sveriges National Atlas. Växter & djur. SNA, Italy.
Wikars L-O (2003) Raggbocken (Tragosoma depsarium) gynnas tillfälligt av hyggen
men behöver gammelskogen. Entomologisk tidskrift 124, 1-12.
Økland B, Bakke A, Hågvar S & Kvamme T (1996) What factors influence the diversity
of saproxylic beeltes? A multiscaled study from a spruce forest in southern Norway.
Biodiversity and Conservation 5, 75-100.
19
Appendix A
Yxnö
Långholmen
Grytsholm
Hästö
Törnsholm
x
x
x
x
x
x
x
x
x
NT
x
x
x
2529 Anthaxia similis
NT
x
x
2646 Ptinus sexpunctatus
VU
x
2695 Stagetus borealis
NT
x
2715 Grynocharis oblonga
VU
2735 Aplocnemus impressus
NT
2753 Ebaeus lapplandicus
VU
2865 Ipidia binotata
VU
3092 Mycetina cruciata
NT
x
x
3534 Prionus coriarius
NT
x
x
4359 Dryophthorus corticalis
NT
x
TC
NT
2425 Stenagostus rufus
VU
x
2468 Ampedus praeustus
NT
2518 Buprestis haemorrhoidalis
NT
2520 Buprestis octoguttata
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Yxnerum
x
Gröskär
Fågelö
Törnsholm
Hästö
Grytsholm
x
Långholmen
x
Yxnö
x
345 Dromius agilis
x
x
x
351 Dromius spilotus
356 Dromius notatus
x
x
Ugglö
Species
x
x
x
Sanningsholmen
Cat nr
x
x
x
Röviken
682 Paromalus parallelepipedus
x
x
x
859 Agathidium rotundatum
867 Agathidium seminulum
Yxnerum
Ugglö
x
Species
Gröskär
Sanningsholmen
x
Cat nr
Fågelö
Röviken
Species dependent on coniferous trees caught in 10 sites in the archipelago of S:t Anna and a
reference site, Yxnerum, on the mainland. Catalogisation was made after Catalogus
Coleopterorum Sueciae (Lundberg 1995). The first table show species found categorized as
red-listed according to Artdatabanken (2003).
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
868 Agathidium laevigatum
996 Gabrius splendidulus
x
1109 Quedius mesomelinus
x
1110 Quedius maurus
1120 Quedius tenellus
x
1334 Bibloporus bicolor
x
1335 Bibloporus minutus
x
x
1352 Euplectus punctatus
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
1354 Euplectus karsteni
x
x
x
1413 Acrulia inflata
x
x
1448 Phloeonomus monilicornis
x
1451 Phloeonomus pusillus
x
1452 Phloeonomus sjoebergi
x
x
x
x
x
x
1600 Phloeocharis subtilissima
1602 Trichophya pilicornis
x
1639 Lordithon lunulatus
x
x
x
x
x
1500 Scaphidium quadrimaculatum
1501 Scaphisoma agaricinum
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
1795 Haploglossa villosula
x
1812 Phloeopora testacea
x
x
x
x
x
x
1873 Dadobia immersa
x
1991 Atheta incognita
x
x
x
x
x
x
2016 Atheta nigricornis
x
x
x
x
2017 Atheta harwoodi
2135 Anomognathus cuspidatus
x
2143 Placusa depressa
x
x
x
x
x
x
x
x
x
2144 Placusa tachyporoides
x
x
x
x
x
x
x
x
2148 Placusa atrata
x
2158 Holobus apicatus
x
x
2159 Holobus flavivornis
x
2344 Dictyoptera aurora
2345 Pyropterus nigroruber
x
x
x
x
x
x
x
x
x
x
x
2394 Malthodes fuscus
2396 Malthodes guttifer
x
2400 Malthodes spathifer
x
x
x
2401 Malthodes crassicornis
x
x
x
2402 Malthodes brevicollis
x
2420 Athous vittatus
2421 Athous haemorrhoidalis
x
x
x
x
2422 Athous subfuscus
x
x
x
x
2423 Limonius aeoniger
x
x
x
x
2442 Prosternon tessellatum
x
x
x
x
2456 Ampedus sanguineus
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2461 Ampedus pomorum
2466 Ampedus balteatus
x
x
1646 Sepedophilus testaceus
2145 Placusa incompleta
x
x
1645 Sepedophilus littoreus
2009 Atheta crassicornis
Yxnerum
Gröskär
Fågelö
Törnsholm
Hästö
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2477 Melanotus villosus
Långholmen
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2478 Melanotus castanipes
x
x
x
x
x
x
2488 Cardiaphorus ruficollis
x
x
x
x
x
x
2517 Buprestis rustica
x
2527 Melanophila cyanea
x
2530 Anthaxia quadripunctata
x
x
x
x
x
x
2532 Chrysobothris chrysostigma
x
2600 Megatoma undata
2645 Ptinus dubius
x
2653 Ptinus subpilosus
x
x
x
x
x
x
2662 Ernobius nigrinus
x
2664 Ernobius mollis
x
2666 Ernobius abietinus
x
x
2667 Ernobius abietis
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2677 Anobium thomsoni
x
x
x
2668 Ernobius pini
x
x
x
x
x
x
x
x
2707 Hylecoetus dermestoides
x
2712 Ostoma ferruginea
x
x
2713 Thymalus limbatus
2724 Thanasimus formicarius
x
x
2658 Dryophilus pusillus
2679 Hadrobregmus pertinax
Yxnerum
x
x
Hästö
x
Grytsholm
x
Yxnö
x
Ugglö
Gröskär
2472 Ampedus nigrinus
Fågelö
x
Törnsholm
2470 Ampedus tristis
Sanningsholmen
Species
Röviken
Cat nr
x
2725 Thanasimus femoralis
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2736 Aplocnemus nigricornis
x
x
x
x
x
x
x
x
x
2742 Dasytes niger
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2760 Malachius bipustulatus
2792 Epuraea boreella
x
x
x
x
2794 Epuraea marseuli
x
x
2803 Epuraea biguttata
x
x
2876 Glischrochilus quadripunctatus
x
2877 Pityophagus ferrugineus
x
x
x
x
x
x
2879 Sphindus dubius
x
x
x
x
x
x
x
x
x
x
x
2880 Arpidiphorus orbiculatus
x
x
x
x
x
x
x
x
x
x
2884 Rhizophagus depressus
x
x
x
x
x
x
x
x
x
2885 Rhizophagus ferrugineus
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2889 Rhizophagus dispar
x
2890 Rhizophagus bipustulatus
x
x
x
x
2911 Silvanoprus fagi
2926 Cryptolestes abietis
x
x
x
2928 Cryptolestes corticinus
x
x
x
2969 Cryptophagus dorsalis
x
x
2993 Atomaria ornata
x
x
x
x
x
x
x
x
x
x
3041 Atomaria bella
x
x
3043 Atomaria pulchra
3051 Triplax russica
x
x
x
x
x
x
x
x
x
x
x
x
3078 Cerylon histeroides
x
x
x
x
x
x
3128 Chilocorus bipustulatus
x
x
x
x
x
x
x
x
x
x
x
x
x
3137 Myzia oblongoguttata
x
x
x
x
x
3138 Myrrha octodecimguttata
3180 Latridius minutus
x
3189 Enicmus rugosus
x
x
3190 Enicmus testaceus
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
3242 Cortinicaria gibbosa
x
x
x
x
x
x
3243 Corticarina similata
x
x
x
x
x
x
3247 Corticarina fuscula
x
x
x
x
x
x
x
3260 Cis alter
x
x
3213 Cartodere constricta
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
3264 Cis comptus
x
3265 Cis hispidus
x
x
3268 Cis boleti
3271 Cis punctulatus
x
x
x
x
x
3274 Cis dentatus
3276 Ennearthron cornutum
x
3285 Hadreule elongatula
x
x
x
x
x
x
x
x
x
x
3298 Bitoma crenata
x
x
3303 Litargus connexus
3319 Chrysanthia viridissima
x
3320 Chrysanthia nigricornis
x
3325 Calopus serraticornis
x
3348 Sphaeriestes castaneus
x
3361 Anidorus nigrinus
x
3425 Uloma rufa
x
3443 Corticeus linearis
x
x
3116 Scymnus limbatus
3130 Exochomus quadripustulatus
x
x
3055 Dacne bipustulata
3114 Scymnus suturalis
Yxnerum
Gröskär
Fågelö
Törnsholm
x
x
3019 Atomaria turgida
Hästö
x
2944 Cryptophagus abietis
2947 Cryptophagus angustus
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
3464 Anaspis frontalis
3466 Anaspis marginicollis
3467 Anaspis thoracica
x
3471 Anaspis rufilabris
x
3472 Anaspis flava
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
3523 Serropalpus barbatus
3536 Spondylis buprestoides
x
x
x
x
x
x
3538 Arhopalus rusticus
x
x
x
x
x
x
3540 Asemum striatum
x
3548 Rhagium inquisitor
x
3564 Alosterna tabacicolor
x
3568 Anoplodera maculicornis
x
x
x
x
x
x
x
x
x
x
x
3571 Anoplodera sanguinolenta
x
x
3580 Leptura melanura
x
x
3569 Anoplodera rubra
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
3598 Hylotrupes bajulus
3631 Pogonochaerus fasciculatus
x
x
x
3638 Acanthocinus aedilis
x
x
x
3750 Cryptocephalus labiatus
x
x
3980 Cimberis attelaboides
x
4145 Brachyderes incanus
x
x
x
4149 Strophosoma capitatum
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
4281 Anthonomus phyllocola
x
x
x
x
x
x
x
x
x
x
4365 Rhyncolus ater
4367 Rhyncolus sculpturatus
x
x
x
x
x
4373 Magdalis phlegmatica
x
x
x
x
x
4376 Magdalis duplicata
x
x
x
x
x
4377 Magdalis frontalis
x
4388 Hylobius abietis
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
4389 Hylobius pinastri
x
x
x
x
x
x
4394 Pissodes castaneus
4395 Pissodes pini
x
x
x
4284 Brachonyx pineti
x
x
x
3506 Hallomenus binotatus
3516 Abdera triguttata
x
Yxnerum
x
x
Gröskär
x
x
Fågelö
x
x
Törnsholm
x
x
Hästö
x
Grytsholm
x
Långholmen
x
Yxnö
Ugglö
3450 Pseudocistela ceramboides
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
x
x
x
x
4397 Pissodes validirostris
x
x
x
x
x
x
x
4399 Pissodes piniphilus
4520 Hylurgops palliatus
x
x
x
x
x
x
x
x
x
x
x
4522 Hylastas brunneus
x
x
x
x
x
x
x
x
x
x
x
4523 Hylastes cunicularius
x
x
x
x
x
Röviken
Sanningsholmen
Ugglö
Yxnö
Långholmen
Grytsholm
Hästö
Törnsholm
Fågelö
Gröskär
Yxnerum
4524 Hylastes attenuatus
x
x
x
x
x
x
x
x
x
x
x
4526 Hylastes opacus
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Cat nr
Species
x
4534 Tomicus minor
4535 Tomicus piniperda
x
4554 Pityogenes chalcographus
x
4558 Pityogenes trepanatus
x
x
x
x
x
x
x
x
4559 Pityogenes quadridens
4560 Pityogenes bidentatus
x
x
4538 Phloeotribus spinulosus
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
4565 Orthotomicus laricis
x
x
x
x
x
x
x
x
4577 Dryocoetes hectographus
x
4580 Crypturgus pusillus
4591 Xyleborus saxesenii
x
x
x
4599 Cryphalus abietis
x
x
x
4601 Pityophtorus micrographus
x
x
4562 Orthotomicus proximus
4576 Dryocoetes autographus
x
x
x
x
x
x
x
x
x
x
x
x
Appendix B
954 Microscydmus nanus
NT
x
1430 Hapalaraea pygmaea
NT
x
1938 Atheta scapularis
NT
2307 Liocola marmorata
VU
x
2597 Globicornis emarginata
NT
x
2598 Globicornis corticalis
VU
x
2686 Xyletinus ater
NT
3053 Triplax rufipes
NT
3307 Mycetophagus piceus
NT
3360 Euglenes oculatus
NT
3424 Uloma culinaris
NT
3448 Prionychus melanarius
EN
x
x
x
3454 Mycetochara axillaris
NT
x
x
x
3461 Scraptia fuscula
NT
x
x
x
3497 Mordellistena variegata
NT
x
3515 Abdera flexuosa
NT
3955 Cassida vibex
NT
x
x
Yxnerum
Gröskär
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Gröskär
Yxnerum
Fågelö
Törnsholm
Yxnö
Ugglö
x
x
x
x
x
96 Bembidion lampros
x
x
x
178 Calathus micropterus
x
x
215 Amara plebeja
216 Amara similata
x
223 Amara lunicollis
x
287 Harpalus rufipes
x
x
83 Trechus quadristriatus
248 Amara majuscula
Fågelö
x
x
x
14 Notiophilus germinyi
230 Amara familiaris
Törnsholm
Hästö
Grytsholm
x
x
x
Sanningsholmen
Species
x
Röviken
Cat nr
Långholmen
x
Hästö
NT
Grytsholm
700 Platysoma minus
x
Långholmen
NT
Yxnö
TC
Ugglö
Species
653 Plegaderus caesus
Sanningsholm
Cat nr
Röviken
Species dependent on deciduous trees caught in 10 sites in the archipelago of S:t Anna and a
reference site, Yxnerum, on the mainland. Catalogisation was made after Catalogus
Coleopterorum Sueciae (Lundberg 1995). The first table show species found categorized as
red-listed according to Artdatabanken (2003).
x
x
x
x
x
x
x
x
x
x
x
x
350 Dromius quadrimaculatus
x
x
413 Hydroporus nigrita
x
560 Helophorus brevipalpis
x
x
x
x
608 Hydrobius fuscipes
x
624 Cercyon lateralis
x
x
630 Cercyon pygmaeus
x
x
x
x
643 Sphaeridium lunatum
x
645 Sphaerites glabratus
x
675 Gnathoncus buyssoni
x
677 Myrmetes paykulli
x
679 Dendrophilus pygmaus
x
x
x
681 Paromalus flavicornis
x
698 Atholus duodecimstriatus
x
736 Ptenidium pusillum
x
x
737 Ptenidium nitidum
774 Acrotrichis sp.
Yxnerum
Gröskär
x
290 Harpalus affinis
639 Cryptopleurum crenatum
Fågelö
Törnsholm
Hästö
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
779 Acrotrichis dispar
x
789 Acrotrichis intermedia
x
x
791 Acrotrichis atomaria
x
794 Acrotrichis sitkaensis
x
x
796 Acrotrichis fascicularis
x
820 Leiodes polita
842 Colenis immunda
x
844 Anisotoma humeralis
x
845 Anisotoma axillaris
x
846 Anisotoma castanea
x
847 Anisotoma glabra
x
x
x
881 Colon brunneum
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
908 Sciodrepoides watsoni
939 Nevraphes angulatus
x
948 Stenichnus scutellaris
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
950 Stenichnus godarti
x
x
848 Anisotoma orbicularis
879 Colon dentipes
x
x
x
951 Stenichnus collaris
x
952 Stenichnus bicolor
x
956 Euconnus claviger
x
973 Nicrophorus vespilloides
x
x
x
x
x
x
x
967 Scydmaenus hellwigii
x
x
x
Gröskär
x
x
x
981 Oiceoptoma thoracica
995 Gabrius coxalus
x
1006 Gabrius trossulus
x
x
1023 Philonthus politus
x
1024 Philonthus succicola
x
1034 Philonthus nigriventris
x
x
1041 Philonthus varians
x
1041 Philonthus varians el confinis
x
1055 Philonthus debilis
1056 Philonthus splendens
x
1077 Platydracus stercorarius
x
x
x
x
x
1111 Quedius cruentus
x
1121 Quedius scitus
x
1122 Quedius xanthopus
1132 Quedius scintillans
x
x
x
1149 Quedius fulvicollis
x
1166 Gyrohypnus atratus
x
1190 Astenus pulchellus
x
1195 Rugilus rufipes
x
x
1196 Rugilus orbiculatus
x
1246 Stenus juno
x
1252 Stenus lustrator
1340 Bibloplectus ambiguus
x
x
1343 Euplectus nanus
1345 Euplectus piceus
x
x
x
1370 Bryaxis bulbifer
x
x
x
x
x
x
x
x
1380 Brachygluta fossulata
x
x
x
x
x
1436 Omalium rivulare
x
x
1444 Omalium caesum
x
x
x
x
x
x
x
1453 Phloeonomus punctipennis
x
x
1480 Acidota crenata
1510 Coprophilus striatulum
x
1530 Carpelimus corticinus
x
1549 Anotylus rugosus
x
x
x
x
x
1608 Mycetoporus lepidus
x
x
1609 Mycetoporus longulus
1610 Mycetoporus bimaculatus
x
x
1393 Megarthrus depressus
1456 Xylodromus brunneipennis
x
x
1375 Tychus niger
1388 Tyrus mucronatus
Yxnerum
Fågelö
Törnsholm
Hästö
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
x
x
x
x
x
x
Yxnerum
Gröskär
Fågelö
Törnsholm
Hästö
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
1635 Lordithon thoracicus
1636 Lordithon exoletus
x
1637 Lordithon trinotatus
x
x
1653 Sepedophilus pedicularius
1655 Tachyporus nitidulus
x
x
1656 Tachyporus obscurellus
x
1662 Tachyporus hypnorum
x
1663 Tachyporus chrysomelinus
x
x
x
x
x
x
1666 Tachyporus atriceps
x
1672 Tachyporus corpulentus
1680 Alaeocharini
x
1682 Tachinus subterraneus
x
x
x
1688 Tachinus marginellus
1700 Aleochara sparsa
x
x
x
x
x
1709 Aleochara fumata
x
1726 Oxypoda opaca
x
1736 Oxypoda umbrata
x
x
1757 Oxypoda formiceticola
x
1776 Calodera aethiops
1814 Phloeopora corticalis
x
x
x
x
x
x
1832 Ischnopoda atra
1840 Dacrila fallax
x
1862 Aleconota gregaria
x
x
x
1869 Liogluta alpestris
x
1880 Philhygra elongatula
x
1890 Philhygra malleus
x
1895 Atheta sp
x
1922 Atheta zosterae
x
1932 Atheta orbata
x
x
1934 Atheta fungi
x
x
x
x
1949 Atheta flavipes
x
1955 Atheta longicornis
x
1978 Atheta setigera
x
2013 Atheta euryptera
x
x
x
x
x
x
x
x
2045 Lymprocorrhe anceps
x
x
x
2022 Atheta brunnea
2023 Atheta nigritula
x
x
x
1939 Atheta sodalis
2021 Atheta nigrifrons
x
x
1815 Phloeopora concolor
1940 Atheta gagatina
x
x
x
x
2064 Amischa analis
x
x
x
x
x
x
x
x
x
x
2162 Oligota inflata
x
x
x
x
x
x
2180 Myllaena intermedia
x
2192 Clambus punctulum
x
2201 Microchara testacea
x
2205 Cyphon ochraceus
2206 Cyphon variabilis
x
2210 Cyphon padi
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2211 Cyphon hilaris
x
2245 Aphodius brevis
x
2247 Aphodius rufipes
x
x
2248 Aphodius depressus
x
2273 Aphodius borealis
x
2294 Serica brunnea
x
x
2306 Cetonia aurata
2312 Trichius fasciatus
x
2316 Platycerus caraboides
x
2350 Lampyris noctiluca
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2354 Podabrus alpinus
2360 Cantharis obscura
Yxnerum
Gröskär
x
x
2091 Lomechusa emarginata
2272 Aphodius ater
Fågelö
x
x
2083 Zyras funestus
2097 Gyrophaena affinis
Törnsholm
Hästö
x
2063 Amischa nigrofusca
2086 Zyras lugens
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
x
x
x
x
x
x
x
2362 Cantharis pellucida
x
2364 Cantharis figurata
x
2365 Cantharis decipiens
x
2373 Cantharis lateralis
x
2375 Rhagonycha fulva
2377 Rhagonycha limbata
x
x
2378 Rhagonycha lignosa
x
x
x
x
x
x
x
2380 Rhagonycha atra
x
2400 Malthodes spathifer
2428 Denticollis linearis
x
2430 Cidnopus pilosus
x
2445 Selatosomus impressus
x
2446 Selatosomus nigricornis
x
2449 Selatosomus aeneus
x
2476 Sericus brunneus
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Gröskär
Yxnerum
Törnsholm
x
Fågelö
Hästö
x
x
x
x
2484 Ectinus aterrimus
2485 Dalopius marginatus
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
x
2499 Microrhagus pygmaeus
x
2510 Aulonothroscus laticollis
2510 Aulonothroscus brevicollis
x
2511 Trixagus dermestoides
x
2512 Trixagus carinifrons
x
x
x
x
x
x
x
x
x
x
x
x
x
x
2602 Ctesias serra
x
x
x
x
2647 Ptinus rufipes
x
2650 Ptinus fur
2656 Hedobia imperialis
x
x
x
x
x
2659 Xestobium rufovillosum
2699 Dorcatoma chrysomelina
x
2703 Dorcatoma dresdensis
x
x
x
2744 Dasytes cyaneus
x
2746 Dasytes aerosus
2747 Dasytes plumbeus
x
x
x
x
x
2868 Thalycra fervida
x
2875 Glischrochilus hortensis
x
2943 Cryptophagus villosus
x
x
x
x
x
x
x
x
2967 Cryptophagus dentatus
x
x
x
x
x
2952 Cryptophagus populi
x
x
2951 Cryptophagus badius
x
x
x
x
x
2972 Cryptophagus scanicus
x
x
x
2989 Atomaria sp.
x
x
x
x
x
3021 Atomaria ruficornis
x
x
x
x
x
x
3042 Atomaria wollastoni
x
x
3050 Triplax aenea
3079 Cerylon ferrugineum
x
x
2827 Meligethes aeneus
3049 Tritoma bipustulata
x
x
2823 Meligethes subaeneus
3031 Atomaria nigrirostris
x
x
x
2820 Meligethes subrugosus
3001 Atomaria fuscata
x
x
x
2817 Meligethes denticulatus
2978 Cryptophagus pilosus
x
x
2573 Dermestes murinus
2606 Anthrenus museorum
x
x
2536 Cytilus sericeus
2540 Agrilus sulcicollis
x
x
x
x
3104 Scymnus frontalis el mimulus
x
3108 Scymnus femoralis
x
3115 Scymnus haemorrhoidalis
x
x
Fågelö
Törnsholm
Hästö
Grytsholm
Långholmen
x
x
x
x
x
x
x
x
x
x
x
x
3135 Tytthaspis sedecimpunctata
x
x
x
3141 Calvia quatuordecimguttata
x
3143 Halyzia sedecimguttata
3144 Psyllobora vigintiduopunctata
x
x
x
x
x
3155 Coccinella magnifica
x
3169 Orthoperus atomus
x
x
3174 Sericoderus lateralis
x
x
x
x
x
x
x
3210 Aridius nodifer
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
3216 Corticaria umbilicata
x
x
x
x
3233 Corticaria longicollis
x
3257 Byturus tomentosum
x
3261 Cis jacquemartii
x
x
3280 Orthocis alni
x
3283 Orthocis vestitus
x
3284 Orthocis festivus
x
x
x
x
x
3354 Salpingus ruficollis
3359 Euglenes pygmaeus
x
3360 Euglenes oculatus/pygmaeus
x
x
x
x
x
x
x
3387 Bolitophagus reticulatus
x
x
x
x
x
3391 Myrmechixenus subterraneus
x
3392 Myrmechixenus vaporariorum
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
3407 Alphitophagus bifasciatus
3411 Palorus depressus
x
x
x
3390 Diaperis boleti
x
x
3203 Stephostethus pandellei
3330 Oedemera flavipes
x
x
x
3202 Latridius angusticollis
3272 Cis fagi
x
x
3192 Enicmus transversus
3230 Corticaria rubripes
x
x
x
3177 Latridius hirtus
x
x
x
3154 Coccinella septempunctata
3193 Enicmus histrio
x
x
x
3126 Platynaspis luteorubra
3139 Propylea quatuordecimpunctata
x
x
x
3104 Scymnus frontalis
3132 Coccinula quattuordecimpustulata
Yxnö
Ugglö
x
Yxnerum
x
Gröskär
3093 Endomychus coccineus
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
x
x
x
x
x
x
x
x
3460 Lagria hirta
x
3474 Tomoxia bucephala
x
3476 Mordella heutheri
x
3482 Mordellistena-sp
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
3485 Mordellistena carinthiaca
x
x
3547 Rhagium mordax
x
x
x
x
x
x
3581 Leptura nigra
3610 Xylotrechus rusticus
x
x
3457 Mycetochara linearis
3578 Leptura quadrifasciata
x
x
3453 Mycetochara flavipes
3499 Mordellochroa abdominalis
x
3613 Clytus arietis
x
x
x
3615 Plagionotus arcuatus
x
3628 Pogonochaerus hispidus
x
3729 Cryptocephalus bipunctatus
x
x
3736 Melasoma populi
x
3791 Linaeidea aenea
3809 Galerucella lineola
x
3810 Galerucella calmariensis
x
3812 Galerucella tenella
x
x
3815 Lochmaea caprea
x
3830 Phyllotreta vittula
x
x
3874 Longitarsus nastutii
x
3886 Altica-sp
x
3893 Altica chamaenerii
x
3898 Batophila rubi
x
3917 Chaetocnema concinna
x
x
x
x
x
x
3924 Chaetocnema hortensis
x
x
3963 Cassida nobilis
3972 Platystomus albinus
3983 Pselaphorhynchites nanus
x
x
x
x
x
x
x
x
x
3986 Pselaphorhynchites germanicus
x
3994 Deporaus mannerheimi
x
x
3995 Deporaus betulae
x
4000 Apion hookeri
4026 Apion fulvipes
Yxnerum
x
3447 Prionychus ater
3452 Isomira murina
Gröskär
Fågelö
Törnsholm
Hästö
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
x
4044 Apion haematodes
x
4048 Apion seniculus
x
x
x
x
x
4091 Otiorhynchus scaber
x
x
4101 Otiorhynchus ovatus
x
x
x
x
x
x
4131 Polydrusus pilosus
x
x
4138 Brachysomus echinatus
x
4148 Strophosoma melanogrammum
x
x
4208 Hypera nigrirostris
x
4245 Cionus longicollis
x
4263 Sibina viscariae
x
4275 Anthonomus pedicularius
x
x
4279 Anthonomus rubi
x
4286 Curculio venosus
x
x
x
x
x
x
x
4295 Rhynchaenus quercus
x
x
x
4297 Rhynchaenus iota
x
4305 Rhynchaenus stigma
4310 Rhampus pulicarius
x
x
4313 Gymnetron pascuorum
x
4329 Miarus campanulae
x
x
4331 Miarus graminis
x
4332 Anoplus plantaris
x
4364 Rhyncolus elongatus
x
4384 Magdalis ruficornis
x
4432 Rhinoncus bruchoides
4443 Coeliodes rubicundus
x
4445 Coeliodes quercus
x
x
x
4521 Hylastes ater
x
4532 Xylechinus pilosus
x
4548 Scolytus ratzeburgi
4551 Scolytus intricatus
4586 Xyleborus dispar
x
x
4258 Tychius picirostris
4296 Rhynchaenus signifer
x
x
4130 Polydrusus cervinus
4292 Curculio pyrrhoceras
x
x
x
4093 Otiorhynchus singularis
4167 Sitona linaetus
x
x
4072 Apion viciae
4128 Polydrusus flavipes
Yxnerum
Fågelö
Törnsholm
x
Gröskär
x
4051 Apion simile
Hästö
Grytsholm
Långholmen
Yxnö
Ugglö
Sanningsholmen
Species
Röviken
Cat nr
x
x
x
x
x
x