Download Alpine Arthropod Diversity

Alpine Arthropod Diversity
Spatial and Environmental Variation
Björn Larsson
Degree project for Master of Science in Biology
Ecological zoology 45 hec
Department of Biological and Environmental Sciences
University of Gothenburg
April 2014
Abstract
Alpine and arctic environments are heavily affected by climate change caused by an ever increasing
emission of greenhouse gasses. Temperatures are estimated to have risen by as much as 3 oC since
preindustrial times. This development threatens this type of habitats as well as all organisms that
inhabit these environments. Knowledge about alpine arthropods is lacking in some areas. Some
groups are better known such as Lepidoptera, Coleoptera and Aranea but even among these there
are clear gaps. This study took place at Latnjajaure field station, located 16 km west of Abisko in
northern Sweden, and looked at the diversity of beetle species as well as the abundance of beetles,
spiders and harvestmen at different altitudes and between two different environments. Samples were
collected using pitfall traps placed every 50 heightmeter at seven altitudes ranging from 1000 to
1300. Eight traps were placed at each altitude, four in open environment and four at or in proximity
to cliffs. An exception was at the 1300 m altitude where only four traps were placed because no
suitable cliff environment was found. The study period was colder than average and had periods of
heavy rainfall which probably had an impact on the results since both temperature and precipitation
seems to have an effect on the activity of the arthropods leading to less individuals and less species
caught. The results of the statistical tests showed that there was no significant difference found in
the diversity of the beetles between either height or environment. There was a significant difference
found between the various altitudes with regards to the number of individuals caught and between
the environments for the spiders and beetles. An interesting find was that there were surprisingly
high numbers of harvestmen found at higher altitudes. They were by far the most numerous of the
studied arthropods. Another interesting find was that there were significantly higher numbers of
arthropods caught at the 1150 m altitude as well as higher numbers of species of beetles found there
compared to the other measured altitudes. These results seem to indicate that there is a shift in
environmental conditions somewhere between the 1150 m altitude and the 1300 m altitude as both
low-alpine and mid-alpine are found there. Among the beetles the most abundant species was
Amara alpina. Most other species were found with only a few individuals. The overall number and
species found were also quite low compared to more southern areas. The results highlights the
importance of further studying alpine environments and the organisms that dwell there in order to
be able to protect them.
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Table of contents
Introduction..........................................................................................................................................3
Background................................................................................................................................3
Aim.............................................................................................................................................5
Materials and methods.........................................................................................................................5
Study site....................................................................................................................................5
Sampling....................................................................................................................................5
Species identification.................................................................................................................7
Analysis......................................................................................................................................7
Results..................................................................................................................................................8
The insect community................................................................................................................8
Statistics...................................................................................................................................13
Discussion..........................................................................................................................................15
Pitfall traps...............................................................................................................................15
Diversity indexes......................................................................................................................15
Weather.....................................................................................................................................15
Evaluation of results.................................................................................................................16
Interesting finds........................................................................................................................18
Conclusions..............................................................................................................................18
Acknowledgements............................................................................................................................19
References..........................................................................................................................................19
Appendix I..........................................................................................................................................23
Appendix II........................................................................................................................................24
Appendix III.......................................................................................................................................25
Appendix IV.......................................................................................................................................29
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Introduction
Background
Alpine and arctic environments are rapidly changing. The main reason for this is anthropogenic
causes due to increased emissions of greenhouse gasses such as carbon dioxide. It is estimated that
the overall global temperature on the planet have risen with about 1 oC since preindustrial times. In
alpine regions however the increase is much greater. Here temperatures might have risen with as
much as 3 oC overall and up to 4 oC during the winter (ACIA, 2005). Effects of this increase is
already visible. Species that previously were only seen occasionally above the treeline are becoming
more common and snowlays are melting out earlier during the summer. This, of course, affects the
species that are adapted to this extreme environment, mainly in the sense that their habitat is
dwindling and that they are being outcompeted by lowland species. Thus, it is important to study
these environments and its inhabitants in order to better understand them and to be able to better
protect them.
There are clear gaps in the knowledge about the diversity of invertebrates in alpine and arctic
environments. Some of the better known groups include Lepidoptera, Coleoptera and Aranaea but
even these groups are not well documented (Nagy et al., 2003). A few previous studies have looked
at arthropods in the study area. For example Brundin (1934) studied the beetle community in the
area surrounding Torne träsk in northern Sweden, where this study took place. Another study looked
at the changes in the insect population between 1998 and 2008 in Padjelanta, also in northern
Sweden (Franzén & Molander, 2011).
Invertebrates have long been used as biological indicators in aquatic ecosystems. A biological
indicator is an organism or a group of organisms that can be used to measure the biotic or abiotic
state of an area due to their reaction to specific changes in the environment (Hodkinson & Jackson,
2005). It has been recognized that different invertebrates have varying degrees of tolerance towards
organic pollution in aquatic ecosystems and that this could be utilized to create effective monitoring
systems. Suggestions have been made that insects might also be useful in assessing changes in
climate in terrestrial ecosystems, particularly mountain ecosystems (Hodkinson & Jackson, 2005). It
has been shown that some insects, for example Neophilaenus lineatus, can vary greatly in their
upper altitudinal limit from one year to the next following the annual mean temperatures (Whittaker
& Tribe, 1996). Carabid beetles have been mentioned as an especially interesting insect group
(Hodkinson & Jackson, 2005). Despite this potential there is no established system for
biomonitoring using invertebrates in alpine and arctic habitats. Therefore it is important to gain
further knowledge about the arthropod community in these ecosystems.
Arthropods in alpine habitats have to be able to endure more extreme environmental conditions than
most arthropods in lower regions (Sømme, 1989). Some factors include high fluctuations in
temperature both during the year and on a daily basis with a possibility of temperatures dropping
below zero during any part of the year. Temperatures also vary more between shaded areas and
more exposed areas. Another differentiating factor is precipitation, a great part of which comes as
snowfall in alpine regions. The amount of precipitation may also vary to a greater extent due to
local conditions such as topology. Furthermore, another factor affecting higher altitudes is increased
wind velocities. Lastly, humidity also varies with altitude since height and temperature affects the
water vapour tension of the air. This leads to a trend in more arid environments with increasing
altitudes. These factors have resulted in a variety of morphological adaptations in alpine arthropods.
One of these adaptations is body size. As a general trend it is possible to see a decrease in size at
higher altitudes (Sømme, 1989). A smaller body size allows insects to take advantage of the
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protected niches in cracks or crevices or under stones (Mani, 1968). Another adaptation is the
tendency for alpine insects to have a darker pigmentation (black, brown or dark red) than lowland
relatives, commonly refered to as melanism. There are several examples of this. One such example
is alpine butterflies of the genus Parnassius in the Himalayans. They have been shown to have
strikingly darker wing markings at 3500 m or higher than at lower altitudes (Mani, 1968). There can
be a number of reasons for this. A darker colour offers additional protection against the inreased
rates of ultra-violet radiation found at higher elevation (Mani, 1968). Being darker also helps
absorbing heat from the sometimes sparse sunlight present and is often more useful as camouflage.
A third trait in alpine arthropods is the loss or reduction of flight wings, referred to as aptery and
brachyptery. A large proportion of species of insects at high altitudes have lost the ability to fly or
have reduced function in the wings (Mani, 1968; Salt, 1954). This may be a response to the increase
in wind velocities found in alpine regions (Sømme, 1989). Flying increases the risk of being carried
away to an unsuitable environment. Dispersal may also be less imortant for insects living in isolated
environments such as high mountain tops due to the fact that so few individuals arrive to replace the
ones that disperse (Nilsson et al., 1993). A further reason may be that the lower temperatures inhibit
a lot of flight activities (Byers, 1969). The fact that many insect species in alpine environments are
very specialized to these extreme conditions makes them vulnarable to rapid large-scale changes in
the climate. They may be considered as an evolutionary “blind alley” (Nagy et al., 2003). This
makes them very interesting from a conservational point of view and further highlights the
importance of studying alpine invertebrate communities.
Most work conducted on alpine beetles are focused on ground beetles (Carabidae). This is probably
beacuse it is a large group with approximately 40 000 species worldwide and can be found in a wide
variety of environments (Lindroth, 1949/1992). They are also often numerous where they are
present and can thus be fairly easily collected compared to other groups. In Fennoscandia there are
about 362 carabid species, 72 of wich occurs in the low alpine zone. Reaching above the low alpine
zone the number of species drastically drops to about 16 (Lindroth 1949/1992). Ecological studies
from the Alps have shown that alpine carabids are “true open land dwellers” and prefer grasslands
or dry stony soils with scattered plant cushions (Brandmayr et al., 2003). They tend to avoid shaded
areas, such as those produced by trees and dwarf-shrubs. They also avoid habitats such as
anthropogenic grasslands, which can be seen as a sign of them being useful as indicators of habitat
condition.
Arachnids are an abundant group in alpine anvironments but as with other invertebrate groups their
species richness decreases with increasing altitude. There are numerous studies on the diversity of
arachnids from the Alps but not so many from other parts of Europe. The most common arachnids
in alpine habitats are spiders and mites. The most common high mountain spiders are epigeal
hunters, ambush predators or web spinners, mostly sheet webs. Most alpine species are diurnal
which is probably due to the colder temperatures at night but some nocturnal species exist.The
activity of alpine arachnids has a peak soon after snow melt and then gradually decreases during the
summer. There are two types of life cycles among alpine arachnids. Stenochronus species that
matures during early summer and has a short adult lifespan and diplochronus that matures later in
the summer and has a longer adult lifespan. Studies in the Alps have shown that some species
exhibit aeronautic behaviour (dispersal by air) in alpine environments. This greatly improves their
ability to disperse across large areas and to colonize new habitats (Thaler, 2003).
Antonsson (2012) studied plant species composition and diversity in cliff ecosystems. His results
showed that cliffs play an important part when assessing alpine landscape diversity. Cliffs maintain
many species that are connected to this type of habitat. He also highlights the importance of
analyzing different species groups seperately because they respond differently in terms of deversity
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patterns. In this study I further investigate differences between open and cliff environments with a
focus on arthropod diversity.
Aims of the study
•
•
•
To find out if there is a change in the diversity, abundance and composition of ground
dwelling arthropods with increasing altitude and between different parts of the season.
To see if it is possible to discern a difference between open habitats and habitats at or in
proximity to cliffs.
To see if precipitation influences the activity of the studied arthropods.
Materials and methods
Study site
The field work took place at Latnjajaure field staion located about 16 km west of Abisko in
northernmost Sweden (coordinates: N7586871, E0643785). It is located at about 1000 m above sea
level (a.s.l.) and the surrounding mountains reach up to 1446 m a.s.l. The mean annual temperature
lies at about -1,4 oC with July being the warmest month with an average temperature of 9,8 oC and
February the coldest with an average temperature of -8,7 oC (Sundqvist, Björk and Molau, 2008).
The valley is covered in snow for the most part of the year and quite large areas are permanently
snowcovered. The landscape in Latnjavagge is very varied with differences in vegetation type over
short distances. These vegetation types include dry heaths, wet meadows, tussoc tundra and screes.
The vegetation is low with small shrubs (Salix spp.) present in some sheltered areas mainly on the
southern slopes. The eastern slopes of the valley are underlain by calcerous materials which is one
of the reasons, along with the diverse landscape, for the high species richness that can be found here
(Lindblad 2007).
The vegetation at the lower altitudes of the study area, up to 1200 m, is fairly similar. The
vegetation type here is a typical mesic meadow with herbs such as Astragalus alpinus, Trollius
europaeus, Ranunculus acris, Bartsia alpina and Poa alpina. At the cliffs we see a different species
composition with Parnassia palustris, Pinguicula alpina, Chamorchis alpina, Thalictrum alpinum
and Carex capillaris present. Cliff ecosystems have been shown to harbour a varied and distinct
community of plant species (Antonsson, 2012). This may be due to the fact that cliffs have a smaller
temperature amplitude as well as higher water availability and higher air humidity (Antonsson,
2012). They tend to be characterised by species that are poor competitors and species that have a
low tolerance for grazing. At 1200 m and above the vegetation type changes to a more dry,
heathlike vegetation with species such as Luzula spicata, Juncus trifidus, Cassiope tetragona and
Festuca ovina dominating. This is consistent with the crossing over from a low-alpine zone into a
mid-alpine zone.
Sampling
The sampling was carried out using pitfall traps. Pitfall traps are small containers (in this case
regular plastic drinking cups) that are sunk into the ground with the brim at the same level as the
ground. In cases where there was not enough soil to allow the trap to be sufficiently sunk the
surrounding soil was raised so that it was at the same level as the trap. Ground dwelling organisms
travelling through the area may fall inte the trap and if constructed correctly they will be unable to
escape (Lövei and Sunderland, 1996). In this study a total of 52 traps were placed at every 50
heightmeter for seven different altitudes ranging from 1000 m to 1300 m a.s.l. On each altitude
eight traps were placed evenly divided between two different environments, open meadows and
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cliffs. The only exception was at 1300 where only four traps were placed due to the difficulty of
finding good places for traps and that no environment corresponding to the cliffs at the lower
altitudes could be found. In order to kill and preserve the captured specimens the traps were filled to
about one fourth with either glycol or a saturated saline solution. In this way the traps could remain
funcional even after quite heavy rainfall. When the traps were emptied the excess conservation
solution was emptied so that the level returned to about one fourth. In cases of extreme rainfall the
excess solution was emptied right after the rain in order to make sure that the traps did not overflow.
The traps were then emptied once every week between the 15th of Juli 2012 and 27th of August
2012. An exception to this is the first sampling that took place irregularly between 26th of June and
5th of July. The samples were then air dried in order for them to be easily transported from
Latnjajaure to the department in Göteborg but were later put in a 70% alcohol solution for storage.
The samples were collected by me during July. During June and August they were collected by Elin
Götmark and Hulda Götmark, who were working at the field station at that time.
Picture 1 shows an example of a pitfall trap. Photo by Ellen Larsson.
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Pitfall trap coordinates
Altitude
N
1000
6821535
1050
6821480
1100
6821504
1150
6821489
1200
6821475
1250
6821433
1300
6821426
E
Meters above sea level
1829765
994
1829929
1045
1830106
1106
1830182
1140
1830396
1207
1830576
1237
1830848
1270
Precision
19
5
4
5
4
3
3
Table 1 shows the coordinates, and their precision, of the pitfall traps. The
coordinates were taken precisely in the middle between the two sets of traps
(environments) at each altitude.
Species identification
During the sampling all organisms that had fallen into the pitfall traps were collected and kept in
seperate containers. The individuals in each sample were sorted, counted and identified using a
stereomicroscope. Among the collected arthropods only the beetles were identified to species level.
The species identification of the beetles was supported by Dr. Thomas Appelqvist (Department of
Biological and Environmental Sciences). The spiders were only devided into the two groups present
namely sheet weavers (Linyphiidae) and wolf spiders (Lycosidae). All other ground dwelling
arthropods collected were counted except for copepods and mites. During the analysis however only
the spiders, harvestmen and beetles were included in the analysis and the total numbers caught.
Analysis
For all analyses the data were entered in and structured using Open Office calc. This program was
also used for the creation of all graphs and tables.
The first analysis was done to see if there was a difference in the diversity of beetles between the
two environments and between the different altitudes. As a measure of diversity two indexes were
chosen, namely the Shannon-Wiener index and Simpson's index. The reason for that two indexes
were chosen was that they have a different approach to abundance within one species. The
Shannon-Wiener index gives a higher priority to rare species while Simpson's index prioritizes more
abundant species. The Shannon-Wiener index measures the uncertainty in predicting the next
species in a given population. One species in the sample gives an index of 0. In theory the index can
get infinitely high but in practice it seldom reaches above 5. Simpson's index measures the
probability that two randomly sampled individuals of an infinately large population are of the same
species and ranges from 0 (one species in the sample) to 1 (Magurran, 2004). The analysis was
devided into two parts. First, a test was done to determine if there was a difference between the
environments. The second test was done in order to determine if there was a difference between the
altitudes. The statistical tests conducted were a Wilcoxon signed ranks test for the first part and a
Kruskall-Wallis ANOVA for the second part. The test for normality used was a KolgomorovSmirnof test. The first test was done in StatView (Abacus Concepts, Inc.) and the second test as
well as the test for normality was done in SPSS (IBM corp., 2013).
A second analysis was conducted to determine if there was a difference in the numbers of
individuals of beetles, harvestmen and spiders caught at different altitudes and between the two
environments. These analyses were done using a Sheirer-Ray-Hare test which is a non-parametric
version of a two-way ANOVA. A non-parametric test was chosen since the test for normality
showed that the data did not conform to a normal distribution (see results). The test for normality
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was done using a Kolgomorov-Smirnof test. All tests conducted for this analysis was done using
SPSS statistics. For the ANOVA, the dependent factor was the variable numbers caught where the
measurments were converted into ranks. The two factors used were height (1000, 1050, 1100, 1150,
1200, 1250 and 1300) and environment (cliff and matrix). The test itself was done in the same way
as a parametric two way ANOVA. The result-table received was then used to manually calculate the
chi2 test statistics with the formula chi2 = factor sum of squares / (corrected total sum of squares /
corrected total df). The value was then compared to a chi2 table to determine the significance
probability. The test method is presented in Statistical and Data Handling Skills in Biology (Ennos,
2012; pp 117-122).
Results
The insect community
In total 3314 individuals of the studied arthropods were collected during the course of the study
period. In addition large amounts of flower visiting diptera and lepidoptera were also caught in the
traps but these were sorted out and not included in the study. Of the studied groups 830 individuals
were spiders, 1873 were harvestmen and 611 were beetles. The 611 beetles were divided among the
rather low number of 34 species (see graph 1). Included among these are two species of
hemipterons (Heteroptera) (Chiloxanthus borealis and Acalypta sp.) and will henceforth be
included in the term beetles. As the graph shows, the beetle community present was completely
dominated by a few species. 75 percent of the total amount of collected beetles belonged to only
eight species. The most common species with 28 percent (174 individuals) of the total finds was
Amara alpina. 62 percent of all the species of beetles collected were found with less than ten
individuals. Among the spiders, 705 were wolf spiders while 125 were sheet weavers. The lower
amount of sheet weavers caught is probably caused by their behaviour. Wolf spiders being more
mobile in their hunt for food than the sheet weavers causing them to come in contact with and get
trapped by the pitfall traps to a greater extent.
Several of the species of beetles caught were alpine specialists restricted to the mountain ranges and
have their main distribution above the treeline. Of the total number of 34 species caught seven were
considered alpine specialists, constituting 20 percent, and an additional four were considered
northern but does have a larger distribution below the treeline as well. They were classified through
looking at the distribution of reports in artportalen (www.artportalen.se). For a list of the alpine
species see table 3. Table 2 shows which species were found at what altitude. As can be seen, the
altitude which had the highest numbers of species was 1150 wich lies in the middle of the range of
altitudes. The altitude with the lowest species count were 1300 shortly followed by 1100. This is
also illustrated by graph 2. This graph also shows tendencies towards an average lower species
density at the cliff environments across all altitudes.
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The total number caught for each species
200
180
160
140
120
100
80
Count
60
40
20
0
Species
Graph 1 shows the number of individuals caught for each species of beeltes.
1000
Amara alpina
Amara quenseli
Anthophagus alpinus
Apion brundini
Atheta sp.
Byrrhus postulatus
Carabus violaceus
Catops luteipes
Chiloxanthus borealis
Cymindis vaporariorum
Eucnecosum brachypterum
Gonioctena arctica
Hypnoides rivularis
Mannerheimia arctica
Miscodera arctica
Mycetoporus punctus
Notiophilus aquaticus
Omalium septentrionis
Patrobus septentrionis
Podabrus lapponicus
Thanatophilus lapponicus
1050
Amara alpina
Acalypta sp.
Anthophagus alpinus
Bryoporus rugipennis
Chiloxanthus borealis
Cymindis vaporariorum
Eucnecosum brachypterum
Hypnoides rivularis
Miscodera arctica
Patrobus septentrionis
Philhygra sp.
Podabrus lapponicus
Quedius fellmani
1100
Amara alpina
Amara quenseli
Anthophagus alpinus
Byrrhus postulatus
Eucnecosum brachypterum
Gonioctena arctica
Miscodera arctica
Omalium septentrionis
Quedius fellmani
Table 2 lists each beetle species found at each altitude.
1150
Amara alpina
Acidota quadrata
Agathidium nigrinum
Anthophagus alpinus
Apion brundini
Atheta sp.
Byrrhus postulatus
Calathus melanocephalus
Catops luteipes
Chiloxanthus borealis
Cymindis vaporariorum
Eucnecosum brachypterum
Gonioctena arctica
Helophorus glacialis
Hypnoides rivularis
Mannerheimia arctica
Miscodera arctica
Notiophilus aquaticus
Olophrum boreale
Omalium septentrionis
Patrobus septentrionis
Podabrus lapponicus
Quedius fellmani
Tachinus elongatus
1200
Amara alpina
Acidota quadrata
Anthophagus alpinus
Atheta sp.
Chiloxanthus borealis
Eucnecosum brachypterum
Gonioctena arctica
Helophorus sibiricus
Miscodera arctica
Notiophilus aquaticus
Olophrum boreale
Patrobus septentrionis
Podabrus lapponicus
Quedius fellmani
Tachinus elongatus
1250
Amara alpina
Acidota crenata
Acidota quadrata
Anthophagus alpinus
Atheta sp.
Bryoporus rugipennis
Byrrhus postulatus
Chiloxanthus borealis
Cymindis vaporariorum
Eucnecosum brachypterum
Latridius minutus
Miscodera arctica
Mycetoporus punctus
Notiophilus aquaticus
Philhygra sp.
Podabrus lapponicus
1300
Amara alpina
Chiloxanthus borealis
Eucnecosum brachypterum
Helophorus glacialis
Helophorus sibiricus
Notiophilus aquaticus
Podabrus lapponicus
Average species caught for each altitude and environment
2,5
2
Count
1,5
Matrix
Cliff
1
0,5
0
1000
1050
1100
1150
1200
1250
1300
Altitude
Graph 2 shows the average number of species of beetles caught for each altitude and
environment.
Alpine
Amara alpina
Helophorus glacialis
Helophorus sibiricus
Manneheimia arctica
Apion brundini
Patrobus septentrionis
Thanatophilus lapponicus
Northern
Anthophagus alpinus
Olophrum boreale
Gonioctena arctica
Quedius fellmani
Table 3 shows the species of beetles that are
considered alpine and northern.
Throughout the study period the total number of specimens collected varied (see graph 3) with the
highest concentrations during the beginning of the sampling period. In the graph, precipitation and
temperature are also represented. The average daily amount of precipitation for each sampling
period showed a good correlation with the concentration of arthropods, with decreased activity with
increased precipitation. The temperature also seemed to be fairly correlated with the concentration
of arthropods caught, though the temperature naturally varies with the amount of precipitation.
Even so, without considering the precipitation it is likely that the decrease in activity during the last
part of the sampling period is caused by shorter days and generally lower temperature.
11
25
12
20
10
8
Count
15
6
10
4
5
2
0
0
1
2
3
4
5
6
7
Precipitation / Temperature
Average numbers caught and precipitation over time
Count
Precipitation
Temperature
8
Sampling
Graph 3 shows the average numbers of arthropods caught per trap, the average precipitation
(given in mm) and the average temperature (given in celcius) at each sampling.
The average numbers of the studied arthropods caught also varied between the altitudes (see graph
4). As can be seen the numbers were highest at 1150 m a.s.l. and lowest at 1100 m a.s.l. Variation
between the environments seemed to be minimal, except at 1150 where it is more visible, and the
variation found is inconsistent between the altitudes. When looking at graph 5 it is clear that the
most common of the studied arthropods were the harvestmen. They were found with higher
numbers at almost all altitudes.
Average numbers caught at each altitude for each environment
25
20
Count
15
Matrix
Cliff
10
5
0
1000
1050
1100
1150
1200
1250
1300
Altitude
Graph 4 shows the average number of catches for each altitude and environment.
12
Average numbers caught for each group at each altitude and environment
14
12
10
Count
8
Beetles
Spiders
Harvestmen
6
4
2
1000
1050
1100
1150
1200
1250
Matrix
Cliff
Matrix
Cliff
Matrix
Cliff
Matrix
Cliff
Matrix
Cliff
Matrix
Cliff
Matrix
0
1300
Altitude / Environment
Graph 5 shows the average numbers caught for each of the studied groups at each altitude
and environment.
Statistics
The Shannon-Wiener index ranged from 1,43 to 2,28 with a mean of 1,80 for the open
environments and 1,68 to 2,43 with a mean of 2,04 for the cliff environments. The values for
Simpson's index ranged from 0,64 to 0,86 with a mean of 0,75 for the open environments and 0,79
to 0,89 with a mean of 0,84 for the cliff environments. The test for normality showed that the data
were significanly different from a normal distribution (see table 3). Both the Shannon-Wiener index
and Simpson's index yielded the same results in this regard. Therefore non-parametric tests were
chosen to calculate the statistics. When observing a barplot of the data the majority of it seemed to
be normally distributed, but it contained too many zeros for a normal distribution to be achievable,
even after transformation. Because of the instability of the data the choice was made to devide the
analysis into two parts in order to have a larger number of replicates for each analysis, thus reducing
the impact of the zeros on the anlysis.
Test of normality, Kolmogorov-Smirnov
Statistic
df
Shannon
0,193
Simpson
0,213
89
89
Sig.
0,000
0,000
Table 4 shows the results of the normality test for the first
analysis.
13
Wilcoxon signed rank test
Statistic
df
Shannon
Simpson
Kruskal-Wallis test
Sig.
Chi-square
df
0,0947
6,430
0,0711
3,540
6
6
Asymp. Sig.
0,377
0,739
Table 5 shows the results of the statistical tests for the first analysis.
The results of the first test showed that there were no significant difference between the cliff
environments and the open environments. The results from the second test showed no significant
difference between the various altitudes (see table 4). For both tests there were no difference in the
results yielded by the two diversity indexes.
For the second analysis the number of individuals caught was used as the variable which reduced
the number of zeros in the dataset. However, the data did not conform to a normal distribution. The
results of the test with the harvestmen showed that there were no significant difference between the
environments but a significant difference was found between the different altitudes. There were no
interaction between environment and altitude. The results for the number of beetles caught showed
that a significant difference was found between both the environments and the altitudes. Again there
were no interaction between the variables. The test for the spiders showed the same results as the
test for the beetles. For the calculated statistics see table 5 and 6.
Test of normality, Kolmogorov-Smirnov
Statistic
df
Harvestmen
0,244
Beetles
0,297
Spiders
0,275
359
359
359
Sig.
0,000
0,000
0,000
Table 6 shows the results of the test for normality for the
second analysis.
Scheirer-Ray-Hare test
Harvestmen
Beetles
Spiders
Environment
Height
Environment*Height
Environment
Height
Environment*Height
Environment
Height
Environment*Height
Type 3 Sum
of Squares
83,225
2863,803
462,146
50,171
246,131
65,854
79,549
732,925
61,548
df
1
6
5
1
6
5
1
6
5
Total mean
Chi-Square Critical value
Square
54,127
1,540
3,841
54,127
52,9
12,592
54,127
8,54
11,07
8,729
5,75
3,841
8,729
28,20
12,592
8,729
7,54
11,07
9,045
8,79
3,841
9,045
81,0
12,592
9,045
6,80
11,07
Table 7 shows the results of the statistical tests for the second analysis.
14
Discussion
Pitfall traps
Pitfall traps are the most commonly used method for measuring the diversity and abundance of a
population of arthropods. This is probably due to the fact that the method is very cheap and labour
efficient. However there are some issues with this sampling method mainly centered around the
assumption that all species are at equal risk of being captured. Since pitfall trapping is a passive
method it relies on the movement of the arthropods for the trapping, more active ones that
frequently travel across larger areas will be caught to a greater extent than ones that are more
stationary. An example of this might be predetors hunting for prey. Topping and Sunderland (1992)
suggested that catches with pitfall traps only represent the relative abundance in a population of
spiders when activity is similar between species which almost certainly in most cases is not true.
Their study also found that differences in vegetation densities surrounding the traps had a species
specific effect on the catch. This might have had an effect on the results when comparing
differences between the two vegetation types studied and also between the different altitudes due to
the more sparse vegetation found at the higher levels.
Different species of carabids have also been shown to have different abilities to escape from pitfall
traps (Halsall and Wratten, 1988; Luff, 1975). Size seems to be an important factor when
determining trapping rate. Notiophilus species for example have been shown to be able to avoid
getting trapped by quickly regaining their balance when encountering a trap or simply spotting the
pitfall by sight and avoiding it (Halsall and Wratten, 1988; Greenslade, 1964). In addition to this a
larger beetle also has an increased risk of being cought in a trap due to the fact that with its
increased speed it can travel across larger areas and therefore encounter more traps (Greenslade,
1964). Some species such as Demetrias atricapillus are very adept climbers which, depending on
the inner surface of the pitfall trap, can increase their ability to excape from the traps (Halsall and
Wratten, 1988). All this may affect the results in the sense that some species may appear more
abundant in relation to other species even though this might not be the case. Though the issues of
pitfall traps have been discussed in several papers there is no real alternative method with the same
effectiveness for sampling arthropods in a reproducible manner.
Diversity indexes
The indexes used to measure the diversity of the beetle community are widly used but there are
some drawbacks with them. Shannon's diversity index assumes an infinitely large population which
of course is impossible. A large population will be less affected by this. It also assumes that all
species in the population have been sampled. This assumption is almost certainly not met due to the
shortcomings of sampling with pitfall traps. There have also been concerns that the index is hard to
interpret since its range is quite small. Simpson's index is interpreted as the probability that two
individuals randomly sampled from a population are of the same species. This interpretation
assumes that the first sampled individual is replaced before sampling another. This assumption is
not met when sampling with pitfall traps. With a large population the effect of this is negligible but
if the population is small the difference can be quite substantial.
Weather
The weather had a major impact on the results of the study. As shown the activity of the arthropods
is very dependent on precipitation and temperature. During the study period there were periods of
heavy rainfall which not only destroyed some traps, as mentioned earlier, but also reduced the
activity of the arthropods resulting in less individuals and thus less species being caught. With less
species and individuals the diversity indexes are of course affected. The study period was also
colder than average which might have had an effect on activity in addition to the rainfall and thus
15
further affect the diversity indexes.
Evaluation of results
From the results the conclusion can be drawn that there is no significant difference in the diversity
of the beetle community between the open environments and the cliff environments. This may be
due to the fact that the arthropods studied are rather mobile, travelling across such a large area in
their search for food that the distance between the localities are not enough to show any difference.
This may also be affected by the use of pitfall traps as the sampling method since they, as
previously mentioned, for example selectively trap species with greater mobility to a larger extent.
The beetles may also be less affected by the different conditions present in the cliff environments
compared to the open environments, which is shown to have an affect on the plant community.
When looking at graph 2, however, there seems to be a tendency for larger amounts of species of
beetles being caught in the open environments at all altitudes. This difference is not as clear at the
lower altitudes but at 1200 and 1250 m a.s.l. it becomes more apparent. It might be beacuse the
environments differ more at higher altitudes in the sense that the cliff environments become more
dry and with more sparse vegetation more rapidly than the open environments which might result in
less occurances of beetles. The statistical tests also showed a slight tendency towards lower species
diversity at the cliff environments but, as mentioned, not significant.
There was also no significant difference found between the altitudes with regards to the beetle
diversity. Looking at graph 2, illustrating the average number of species caught, this pattern, or
rather lack of pattern, can also be seen. These results were quite intreresting as it would be expected
that the average diversity would decrease with increasing altitude due to the harsher climate present
there.
One interesting observation however is the increase in number of species caught at the 1150 m
altitude. Here the number of species caught were markedly (though not significant) higher for both
environments. The sites for the traps were chosen to be as similar as possible, except for the natural
change in vegetation and climate with increased altitude, so the cause for this difference is not
directly obvious. One hypothesis is that the threshold between the low-alpine zone and the midalpine zone is located somewere around the 1150 m altitude. This would result in a higher species
count at this altitude since species specialized for each zone overlap there. This hypothesis has not
been tested however but there are some signs pointing in this direction. Up to 1150 m above sea
level the vegetation is fairly similar but reaching above it slowly becomes more sparse and meagre.
Looking at the beatles caught at this altitude there are some species that were found there and at
altitudes above but not below and species that were found there and below but not above, presented
in table 7. These observations could point towards a shift close to the 1150 m level, but again, this
has not been tested. Similar results were obtained by Antonsson et al. (2009). In their study they
looked at nurse plant effects by the cushion plant Silene acaulis along an altitudinal gradient in the
same area in northern Sweden as this study took place. Their results showed that there was a shift in
the difference between the number of species inside the cushions and outside from slightly negative
at lower altitudes to positive at altitudes above 1300 m. These facts further point twards a shift in
environmental conditions somewhere between 1150 m a.s.l. and 1300 m a.s.l.
There were also two species that were found only at 1150 m contributing to the higher species count
there, namely Agathidium nigrinum and Calathus melanocephalus. Agathidium nigrinum feeds
mainly on slime molds (Mycetozoa) and thus are aggregated to locations were these are present.
Calathus melanocephalus was only caught with one specimen. Thus both these species were
considered chance finds and not connected to the altitude in any particular way.
16
At 1150 m and below
Apion brundini
Catops luteipes
Hypnoides rivularis
Mannerheimia arctica
Omalium septentrionis
At 1150 m and above
Acidota quadrata
Helophorus glacialis
Olophrum boreale
Tachinus elongatus
Table 8 shows species present at 1150 m and above and
at 1150 m and below.
That the statistical tests for the diversity of the beetles gave unsignificant results might be caused by
the overall small numbers of species caught. A lot of the samples did not contain any beetles at all
wich resulted in the dataset containing a lot of zeros, affecting the stability of the dataset. Further
sampling during several seasons may result in better data and more clear results. As previously
mentioned the weather during the sampling was rainier than average with periods of heavy rainfall.
This most certainly affected the catches of the pitfall traps as they were occasionally filled with
water resulting in loss of the caught arthropods as well as preventing the trap from catching
anything else until they were emptied of water. In addition to this the temperatures were also lower
than average which probably also affected the activity of the studied arthropods.
An interesting observation is the surprisingly high numbers of harvestmen caught at all altitudes.
They were by far the most common among the studied arthropods and comprised about 55% of the
total numbers caught. When compared to the numbers of beetles and spiders cought for each
altitude the proportion of harvestmen remains around 50% while the proportion of beetles decreases
somewhat and the proportion of spiders increases with increasing altitude. The exception to this was
at 1300 m a.s.l. where the proportion of harvestmen rises to slightly over 80%. At this altitude the
vegetation changes quite rapidly in response to the change from a lowalpine to a midalpine climate
which could explain the change. The statistical analysis showed some support for there being
generally higher numbers of harvestmen caught at higher altitudes (1150 m and above) compared to
the lower. This may be due to the change in vegetation that occurs at these altitudes, suggesting that
the harvestmen prefer low, sparse vegetation compared to the relatively lush vegetation found at the
lower altitudes. That the harvestmen were caught with a higher frequency at the cliff environment at
the lowest altitudes could offer some support for this.
Yet again the 1150 m altitude stands out with the highest numbers caught for both harvestmen and
beetles at both the cliffs and the open environments. The difference is quite pronounced as well with
aproximately twice as many harvestmen caught in the open environments compared to the altitude
with the second highest numbers. The same is true for the beetles. Only the spiders were caught
with a higher frequency at another altitude. This could further point towards the fact that there is an
unknown factor that differs at this altitude compared to the others since this difference is supported
by the results of the statistical test. The higher number of individuals caught might also be the
cause for the higher number of species of beetles that were found at this altitude. For the spiders
there was no clear pattern in their distribution. There seems to be a slightly higher density at the
higher altitudes, somewhat supported by the statistics. The exception to this was at 1300 wich had
the lowest amount caught of all the altitudes. One possibility for this could be that the spiders prefer
a more sheltered environment which is offerd by the cliffs or by vegetation. At some altitudes,
mainly the higher ones, there is a small but noticable increase in numbers caught at the cliff
environments. It could also be related to the availability of places to escape sudden periods of cold
temperatures. At 1300 m above sea level it is quite common with snow even during the summer.
17
Interesting finds
The most common species of beetle found was, as previously mentioned, Amara alpina with a total
of 174 finds. Constituting almost 30 percent of all individuals collected and found at all sampled
altitudes this species seems well adapted to an alpine habitat. In contrast some of the species of
beetles collected are quite uncommon with only a few finds from the Swedish mountain ranges.
Examples of these include Apion brundini, Helophorus sibiricus and Helophorus glacialis. Some of
these species were also collected the year before this study, again using pitfall traps, as well as
another rare species, Diacheila arctica. That they were found both years and with more than one
individual (all except Diacheila arctica from which only one specimen was caught) might point
towards that these species are actually more frequent than what current data suggests, the lack of
inventories conducted in alpine regions being the most probable cause for this. This further
highlights the importance of continuing to study this environment and its organisms. Two
interesting finds were Apion brundini and Carabus violacius ssp. arcticus that were both found with
few individuals mainly at the lower altitudes. Both species are limited to alpine regions and both are
endemic to the scandinavian mountain ranges. Apion brundini has its closets sister species (Apion
amethystinum) located in areas around the Kaukasus. Both species also lack fully developed flightwings, resulting in quite limited dispersal abilities. This has been seen as an indication that they
survived the latest ice age close to their present habitats which raises the question of how they
survived.
It has long been debated how species inhabiting the cold environments present in Europe's
mountain ranges survived the last ice age. For temperate species in general it has been hypothesised
that they survived at refugia in southern Europe, but for alpine species the answer is not as clear.
There are two main hypotheses that are being discussed (Lohse et.al., 2011). One hypothesis,
referred to as the massif de refuge hypothesis, claim that alpine species survived at refugial areas in
southern and eastern Europe. The other hypothesis holds that these species survived on isolated
icefree areas within the limits of the ice sheet, so called nunataks (Lohse et.al., 2011; Westergaard
et. al., 2011). For a long time the “nunatak theory” was considered the main hypothesis but during
the past 20 years there have been a lot of molecular data presented that support the massif de refuge
theory (Brochmann, 2003; Skrede, 2006). Westergaard et. al. (2011) however presented molecular
data from two rare arctic plants that favours the nunatak theory. Lohse et. al. (2011) studied
molecular data from alpine carabid beetles, genus Trechus, and its local radiation in the alps and
found that a combination of the two theories was the most likely explanation.
Conclusions
As a conclusion it can be said that the beetle divesity is rather low in this alpine habitat compared to
more southern areas. The density of the beetles was also a bit on the low side for most of the
species, Amara alpina excluded. The low density might also have affected the total numbers of
species caught. Sampling over several seasons and in a greater variety of environments would
probably yield a higher species count. Sampling over several seasons also reduces the negative
impact caused by the weather as discussed previously. Another way to increase the species count is
to complement the pitfall traps with other catching methods as the traps can be selective in what
species that get caught. For example pitfall traps would not catch species that spend most of their
time feeding on the branches of shrubs or other higher plants, except by chance. For those species a
catching net would be a better choice.Thus, this study may not provide a good estimate of the true
diversity of the beetle population in an alpine habitat but rather add to the overall knowledge about
the beetle community present.
For further studies it might be interesting to look more closely at the harvestmen. Their abundance
18
in this habitat was rather surprising and the question arises on what they feed at the higher altitudes.
Many harvestmen are predators eating primarily smaller insects, but some feed on plant material or
fungi while some still are scavengers. One possibility is mites, that were noticed to be abundant in
the traps but, as noted in the beginning, were not counted. Still the increase in abundance with
increasing altitude is interesting. Another continuation could be to look at the beetles that are
present mainly around snowlays. Many speceis are depentent on snowlays for foraging or wintering
etc. Snowlays as a habitat is threathened due to climate change resulting in earlier meltout and thus
the species that inhabit this habitat comes under threat (Björk & Molau, 2007). Monitoring them
could be important to preserve their diversity.
19
Acknowledgments
I would like to thank my supervisors Ulf Molau and Thomas Appelqvist for all their support with
project planning, statistics and species identification. Elin Götmark and Hulda Götmark for their
invaluable help with finding locations for the traps and sampling them during June and August. The
Swedish Polar Research Secretariat and Abisko Research Station for use of their facilities. Lastly I
would like to thank everyone else that has participated in or contributed to this master project.
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22
Appendix I: Beetle species list
Species
Acalypta sp.
Acidota crenata
Acidota quadrata
Agathidium nigrinum
Amara alpina
Amara quenseli
Anthophagus alpinus
Apion brundini
Atetha sp.
Bryoporus rugipennis
Byrrhus postulatus
Calathus melanocephalus
Carabus violaceus
Catops luteipes
Chlioxanthus borealis
Cymindis vaporariorum
Eucnecosum brachypterum
Gonioctena arctica
Helophorus glacialis
Helophorus sibiricus
Hypnoides rivularis
Latridius minutus
Mannerheimia arctica
Miscodera arctica
Mycetoporus punctus
Notiophilus aquaticus
Olophrum boreale
Omalium septentrionis
Patrobus septentrionis
Philhygra sp.
Podabrus lapponicus
Quedius fellmani
Tachinus elongatus
Thanatophilus lapponicus
Total individuals caught
Number of species
Altitude and environment (M=Matrix, C=Cliff)
1000M 1000C 1050M 1050C 1100M 1100C 1150M 1150C 1200M 1200C 1250M 1250C 1300M
1
1
1
1
1
1
1
2
62
45
8
18
5
3
1
58
23
8
2
1
1
1
1
8
1
4
12
1
1
3
3
17
28
3
1
1
1
1
2
1
1
1
1
2
1
1
1
1
1
1
1
1
1
1
1
2
4
8
1
4
12
1
2
15
3
3
2
2
3
3
2
3
1
1
1
1
6
17
3
7
1
5
1
1
2
1
4
3
4
1
1
1
1
2
1
4
5
1
1
3
1
1
3
1
2
1
5
1
3
2
1
2
2
2
1
3
1
3
8
3
2
1
6
2
1
5
1
6
3
2
1
1
5
10
1
1
1
2
2
1
1
1
1
1
4
2
1
78
59
37
19
16
14
179
89
39
9
31
13
28
14
16
9
10
6
9
20
15
12
6
14
9
7
Table 1 lists the species of beetles found and shows at which location and with how many
speciemens it was caught.
23
Appendix II: Map over Latnjavagge
Appendix III: Trap stations
Picture 1 shows the open and cliff environments at the 1000 m altitude. Photo by Ulf Molau.
Picture 2 shows the open environment at the 1050 m altitude. Photo by Ulf Molau.
25
Picture 3 shows the cliff environment at the 1050 m altitude. Photo by Ulf Molau.
Picture 4 shows the open and cliff environments at the 1100 m altitude. Photo by Ulf Molau.
26
Picture 5 shows the open and cliff environments at the 1150 m altitude. Photo by Ulf Molau.
Picture 6 shows the open and cliff environments at the 1200 m altitude. Photo by Ulf Molau.
27
Picture 7 shows the open and cliff environments at the 1250 m altitude. Photo by Ulf Molau.
Picture 8 shows the open environment at the 1300 m altitude. Photo by Ulf Molau.
28
Appendix IV: Beetle images
Acalypta sp.
Acidota crenata.
Acidota quadrata.
Agathidium nigrinum.
Amara alpina.
Amara quenseli.
Anthophagus alpinus.
Apion brundinii.
29
Atheta sp.
Bryoporus rugipennis.
Byrrhus postulatus.
Calathus melanocephalus.
Carabus violaceus.
Catops luteipes.
Chiloxanthus borealis.
Cymindis vaporariorum.
30
Eucnecosum brachypterum.
Gonioctena arctica.
Helophorus glacialis.
Helophorus sibiricus.
Hypnoides rivularis.
Latridius minutus.
Mannerheimia arctica.
Miscodera arctica.
31
Mycetoporus punctus.
Notiophilus aquaticus.
Olophrum boreale.
Omalium septentrionis.
Patrobus septentrionis.
Philhygra sp.
Podabrus lapponicus.
Quedius fellmani.
32
Tachinus elongatus.
Thanatophilus lapponicus.
The images in this appendix are not meant for species identification and are not in a comparable
scale.
33