Alpine Arthropod Diversity [PDF]

chosen, namely the Shannon-Wiener index and Simpson's index. The reason for that two indexes were chosen was that they h

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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 3

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 4

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 5

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 7

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.

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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

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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.

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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.

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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.

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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.

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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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Greenslade P.J.M. (1964). Pitfall trapping as a method for studying populations of Carabidae (Coleoptera). Journal of Animal Ecology, 33:301-310. Halsall N.B and Wratten S.D. (1988). The efficiency of pitfall trapping for polyphagous predatory Carabidae. Ecological entomology, 13:293-299. Hodkinson I.D. & Jackson J.K. (2005) Terrestrial and aquatic invertebrates as bioindicators for environmental monitoring, with particular references to mountain ecosystems. Environmental Management, 35:649-666. IBM Corp. Released 2013. IBM SPSS Statistics for Windows, Version 22.0. Armonk, NY: IBM Corp. Lindblad K. (2007). Tundra landscape ecology: diversity across scales (Ph. D. Thesis). Gothenburg, University of Gothenburg. Lindroth C.H. (1949/1992). Ground beetles (Carabidae) of Fennoscandia. New Dehli: Amerind Publishing Co. Pvt. Ltd. [Original titel: Die Fennoskandischen Carabidae]. Lohse K., Nicholls J.A. and Stone G.N. (2011). Inferring the colonization of a mountain rangerefugia vs. nunatak survival in high alpine ground beetles. Molecular Ecology, 20:394-408. Luff M.L. (1975). Some features influencing the efficiency of pitfall traps. Oecologia, 19:345-357. Lövei G.l. & Sunderland K.D. (1996). Ecology and behavior of ground beetles (Coleoptera: Carabidae). Annual Reviews Entomology, 41:231-256. Magurran A.E. (2004). Measuring biological diversity. Oxford, Backwell publishing. Mani M.S. (1968). Ecology and Biogeography of High Altitude Insects. Series Entomologica vol 4. Junk, The Hague, 527 pp. Mikhailov Y.E. and Olschwang V.N. (2003). High altitude invertebrate diversity in the Ural Mountains. In: Nagy L., Grabherr G., Körner Ch. and Thompson D.B.A (Eds.), Alpine diversity in Europe. Ecological studies, 167:259-279. Nagy L., Grabherr G., Körner Ch., Thompson D.B.A. (2003). Alpine Biodiversity in Europe. Ecological studies vol 167. Springer-Verlag Berlin Heidelberg, 477 pp. Nilsson A.N., Petterson R.B. and Lemdahl G. (1993). Macroptery in altitudinal specialists versus brachyptery in generalists – a paradox of alpine scandinavian carabid beetles (Coleoptera: Carabidae). Journal of Biogeography, 20:227-234. R Core Team (2012). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0, URL http://www.R-project.org/ Salt G. (1954). A contribution to the ecology of Upper Kilimanjaro. Journal of Ecology, 42:37 5423.

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Skrede I., Eidesen P.B., Portela R.P. and Brochmann C. (2006). Refugia, differentiation and postglacial migration in arctic-alpine Eurasia, exemplified by the mountain avens (Dryas octopetala L.). Molecular Ecology, 15:1827–1840. Sømmme L. (1989). Adaptations of terrestrial arthropods to the alpine environment. Biological Reviews, 64:367-407. Thaler K. (2003). Diversity of high altitude Arachnids (Araneae, Opiliones, Pseudoscorpiones) in the Alps. In: Nagy L., Grabherr G., Körner Ch. and Thompson D.B.A (Eds.), Alpine diversity in Europe. Ecological studies, 167:281-296. Topping C.J. and Sunderland K.D. (1992). Limitations to the use of pitall traps in ecological studies exemplified by a study of spiders in a field of winter wheat. Journal of Applied Ecology, 29:485491. Westergaard K.B., Alsos I.G., Popp M., Engelskjøn T., Flatberg K.I. and Brochmann C. (2011). Glacial survival may matter after all: nunatak signatures in rare European populations of two westarctic species. Molecular Ecology, 20:376-393. Whittaker J.B. & Tribe N.P. (1996). An altitudinal transect as an indicator of responses of a spittlebug (Auchenorrhyncha: Cercopidae) to climate change. European Journal of Entomology, 93:319–324.

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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.

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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.

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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.

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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.

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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.

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Appendix IV: Beetle images

Acalypta sp.

Acidota crenata.

Acidota quadrata.

Agathidium nigrinum.

Amara alpina.

Amara quenseli.

Anthophagus alpinus.

Apion brundinii.

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Atheta sp.

Bryoporus rugipennis.

Byrrhus postulatus.

Calathus melanocephalus.

Carabus violaceus.

Catops luteipes.

Chiloxanthus borealis.

Cymindis vaporariorum.

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Eucnecosum brachypterum.

Gonioctena arctica.

Helophorus glacialis.

Helophorus sibiricus.

Hypnoides rivularis.

Latridius minutus.

Mannerheimia arctica.

Miscodera arctica.

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Mycetoporus punctus.

Notiophilus aquaticus.

Olophrum boreale.

Omalium septentrionis.

Patrobus septentrionis.

Philhygra sp.

Podabrus lapponicus.

Quedius fellmani.

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Tachinus elongatus.

Thanatophilus lapponicus.

The images in this appendix are not meant for species identification and are not in a comparable scale.

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