Fauna norvegica 2019 Vol. 39: 77–110. Environmental conditions limit the distribution of arcticus (, ) in lakes on the mountain plateau, Southern

Tore Qvenild1 and Trygve Hesthagen2

Qvenild T and Hesthagen T. 2019. Environmental conditions limit the distribution of (Branchiopoda, Notostraca) in lakes on the Hardangervidda mountain plateau, Southern Norway.Fauna norvegica 39: 77–110.

The Arctic tadpole shrimp Lepidurus arcticus has a circumpolar distribution where the Hardangervidda mountain plateau in Norway marks its southernmost limit. Within this area, we searched for L. arcticus in 238 lakes in 27 catchments. On Hardangervidda, the distribution pattern of L. arcticus is highly skewed. In the 16 catchments located in the central and eastern parts, L. arcticus was recorded in 70% of all the lakes studied (n=191). The remaining 11 catchments located in western areas, are almost free of lakes with L. arcticus (n=47). The most striking difference between these two areas is the significantly higher level of snow deposition in the western areas. This delays the ice break-up, which results in lower water temperatures and a shorter growing season. The water of lakes in western areas (N=36) is also more dilute than those in the central and eastern areas (N=201), with mean calcium concentrations of 0.81±0.48 and 1.62 ± 1.12 mg L-1, respectively. In the lakes in the central and eastern areas hosting L. arcticus (N=95), the mean value was slightly higher (1.67 ± 1.14 mg L-1). The combination of low water temperature, a short growing season and dilute water low in calcium may explain the near total absence of L. arcticus in the western part of Hardangervidda. All lakes contain brown trout Salmo trutta, and as L. arcticus is heavily sought for as food, the analyses of fish stomachs are the most reliable method of detecting the species. However, this prey-predator relationship may severely reduce the population of L. arcticus, and their presence may also be a function of the proximity of species refugia. This is evident in the context of fish predation, but also of water quality. Hence, in the central and eastern parts of the plateau, where L. arcticus is common, their occurrence increased significantly with lake size, being found in 54% of the lakes <1.0 km2, as opposed to 97% in the bigger lakes. Furthermore, L. arcticus is most frequently found in lakes at altitudes between 1100 and 1299 m a.s.l. We conclude that environmental constraints limit the distribution of L. arcticus on Hardangervidda. The projected increase in temperature towards the end of this century may exterminate L. arcticus from the lower parts of Hardangervidda, especially in the most shallow lakes. Many of the lakes have water quality with pH <6.0 and calcium concentration <1.0 mg L-1. In such lakes L. arcticus is living on the edge of its survival, and the projected increase in precipitation may dilute the waters even further, pushing L. arcticus nearer to its extinction threshold.

doi: 10.5324/fn.v39i0.2687. Received: 2018-08-19. Accepted: 2019-06-05. Published online: 2019-07-11. ISSN: 1891-5396 (electronic).

Keywords: Arctic tadpole shrimp, Salmo trutta, water quality requirements, thermal conditions, climate change

1. County Governor of , Statens hus, Parkgata 36, NO-2317 Hamar, Norway 2. Norwegian Institute for Nature Research (NINA), PO Box 5685, Torgården, NO-7485 Trondheim

Corresponding author: Tore Qvenild E-mail: [email protected]

77 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

INTRODUCTION which L. arcticus are reported is 1.52 mg L-1 (Borgstrøm et al. 1976). However, only three lakes were investigated and hence, The Arctic tadpole shrimp Lepidurus arcticus (Pallas, 1793) has the lower lethal threshold for calcium demand has yet to be a circumpolar distribution and occurs in both shallow moraine established for L. arcticus. Rapid postmoult calcification of the ponds and large lakes in subarctic areas (Hessen et al. 2004). In exoskeleton is essential for all , and lack of calcium its central circumpolar area, L. arcticus is mainly found in ponds may limit their success in localities with such water quality and fishless shallow lakes (Jeppesen et al. 2001; Lakka 2013). (Rukke 2002). However, in Scandinavia it has been described as a boreo-alpine The water temperature is also vital to all aquatic crustaceans. relict species that occurs at sea-level in the Arctic region, and The thermal conditions of lakes are not only a result of summer at progressively greater altitudes towards the south (S. Sømme temperatures, but also related to snow accumulation during the 1934; Økland & Økland 2003). On the Norwegian mainland, it winter. Snow deposition and spring temperature are the main is a typically alpine species, mainly co-existing with fish (S. drivers of ice break-up, and determine the water temperature Sømme 1934; Aass 1969). In ponds and small shallow lakes in regime (Kvambekk & Melvold 2010). In catchments with heavy its southern outreach on the Scandinavian peninsula, which is snowfall, the ice break-up may be delayed and thus lead to a often fishless, L. arcticus seems to be absent (S. Sømme 1934; thermal deficit that impedes the development of L. arcticus Berg 1954; Blomkvist 1995; Klausen 2012). (Qvenild et al. 2018). Snow deposition on Hardangervidda is The Hardangervidda mountain plateau in Southern Norway highly variable, with an almost fourfold increase in an east to is the largest peneplain (eroded plain) in Europe (Anonymous west gradient (Qvenild et al. 2018). 1974). This plateau marks the southernmost outreach of L. In this study we hypothesised that the combination of low arcticus (S. Sømme 1934; Økland & Økland 2003). The first to water temperature, short growing season and dilute water may describe L. arcticus from this mountain plateau was Huitfeldt- impede ecdysis in L. arcticus. Furthermore, this particular Kaas (1911), who offered only a general description of the is a large food item and is heavily fed upon by fish species as being an important food item for brown trout Salmo (Dahl 1920; Aass 1969; Borgstrøm et al. 1985; Jeppesen et al. trutta Linnaeus, 1758. He used the name Apus glacialis (Braem, 2001; Qvenild & Rognerud 2018; Presthus Heggen et al. 2010). 1893) which together with Lepidurus glacialis (Packard, 1883) Species “living on the edge” may be exterminated by reinforced was commonly used in the early decades of the 20th century vectors such as fish predation (Aass 1969; Jeppesen et al. 2001; (Økland & Økland 2003). Other synonym names from this Presthus Heggen et al. 2010). Our key question is whether the period were Monoculus arcticus (Pallas, 1793), Lepidurus distribution pattern of L. arcticus may be explained by these spitzbergensis (Bernard, 1882), Apus productus var. glacialis environmental variables. Furthermore, alpine ecosystems are (Braem, 1893) and Lepidurus ussuriensis (Sidorov, 1927). expected to suffer severely from the effects of global warming No Norwegian name was known for L. arcticus at that time. (Lindholm et al. 2012, 2015) and climate change may be a threat The Norwegian name “skjoldkreps” (“scale shrimp”) was to L. arcticus (Hessen et al. 2004; Lakka 2013; Qvenild et al. introduced by Dahl (1913). This is astonishing as L. arcticus is 2018; Väinölä et al. 2019). Detailed mapping of L. arcticus quite conspicuous due to its large size and archaic appearance, is therefore an essential aspect of identifying future range and it was well known by local fishermen. On the other hand, retractions. the other large crustacean, the amphipod Gammarus lacustris G.O. Sars, 1863, had a variety of Norwegian local names such as “marflo, matflo, grunnåte, nettskjær, etc.” (Dahl 1915). METHODS AND MATERIAL The brown trout on Hardangervidda are of good size and Study area quality and support a number of traditional fisheries (I. Sømme The Hardangervidda landscape is characterized by barren, 1941). Their main crustacean food items are L. arcticus, G. treeless moorland interrupted by numerous pools, lakes, rivers lacustris and Eurycercus lamellatus A.F.M (Huitfeldt-Kaas and streams. The plateau covers an area of about 9800 km2, 1911; Dahl 1917; I. Sømme 1941). Information regarding their of which the most remote areas account for about 8000 km2 occurrence has long been fragmentary and incomplete, and (Anonymous 1974). In this area we included 27 catchments not until the beginnings of the 1970s did a rough distribution that cover 6569 km2 (Figure 1, Table 1). We have divided these pattern appear that included almost all the records from the catchments into two groups. Of the “11 western catchments”, central and eastern parts of the plateau (Anonymous 1974). ten drain to the west, while of the “16 central and eastern The geology of Hardangervidda is highly variable and catchments”, nine drain to the east and seven to the west. The this is reflected in the water chemistry of the lakes, many extent of these catchments was calculated using the NEVINA of whose waters are low in pH and calcium (Skjelkvåle & procedure (NVE Atlas, nve.no). Within this area, we have Henriksen 1998). Lepidurus arcticus is highly sensitive to acid identified 930 named lakes covering 656 km2 with a median water (Borgstrøm & Hendrey 1976; Lakka 2013) becoming size 0.162 km2 (range 0.004 – 78.77 km2). These localities also extinct in waters with a pH below 5.5 (Fjellheim & Raddum included 27 reservoirs for hydropower production ranging in 1990). The lowest ambient calcium concentration in lakes in surface area from 0.48 to 78.77 km2 (Appendix 1). In addition,

78 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019 there are approximately 11,600 unnamed lakes and ponds almost all lakes (Appendix 1). covering 157 km2 (range 0.0006 – 0.7035 km2). Most of the 36 Calcium concentration and pH are vital variables in the natural lakes with bathymetric maps on Hardangervidda have survival of L. arcticus. There are significant intermediary and mean depths <10 m (Appendix 1). Most of the lakes within the seasonal variations in calcium concentration within a lake. The study area are situated at altitudes of 1100 – 1399 m a.s.l. (85%), highest values of calcium are usually measured in the spring while 11% and 4% are located at lower and higher altitudes, when the concentration may be more than twice the minimum respectively. values registered later in the season (Fjellheim et al. 2002, 2007, The western area is dominated by rocky terrain and expanses 2018). This variability is most prominent in small lakes and of bare rock with thin or no moraine-covered bedrock of brooks. In bigger lakes the variation follows the same pattern Precambrian gneisses and granites and sparse or no vegetation. but within a much more limited range. On the other hand, the Some catchments in the central part comprise bedrock of lowest pH values are measured in the run-off period in the Cambro- sedimentary origin. The bedrock in the spring. However, we assume that the low ambient pH-levels eastern areas also covers gneisses and granites but has deeper encountered throughout the growth period in summer is most layers of moraine. The water chemistry on Hardangervidda critical for L. arcticus as the run-off period is normally short closely reflects the local bedrock geology (Skjelkvåle & and their resting eggs are very resistant to external stresses Henriksen 1998). Hence, the lakes are highly variable in (Longhurst 1955; Blomquist 1995; Lakka 2013). Hence, we water chemistry, ranging from lakes being practically free of have used the minimum values of calcium and the concurrent electrolytes to lakes with high ionic strength. The content of pH-value in summer-autumn when water quality series are total organic carbon (TOC) is very low, with <2.0 mg C L-1 in available for assessing the occurrence of L. arcticus. Almost all

Figure 1. Map of the Hardangervidda with the 27 catchments indicated. The boundary of the study area is marked by a blue line. Details of the catchments are shown in Table 1. The study area is divided into two groups with 11 western catchments (W) and 16 central and eastern catchments (C/E). These areas are divided by a dotted line. Norwegian Mapping Authority CC BY 4.0.

79

Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

L. arcticus L. with Proportion (%) of lakes lakes of (%) Proportion 20 0 100 63 100 100 53 100 0 40 0 0 0 0 0 0 0 0 33 69 29

Norwegian arcticus

L. L. with lakes of No.

2 0 1 5 1 3 10 2 0 6 0 0 0 0 0 0 0 0 3 29 2 No. of lakes examined lakes of No.

10 1 1 8 1 3 19 2 1 15 3 4 1 1 20 3 1 2 9 42 7 No. of lakes of No. 22 14 6 22 2 14 37 2 2 31 4 10 1 2 46 13 7 34 44 99 103

). Details). the of specific lakes are given

Lake area km area Lake 2

13 4 2 18 0 9 10 1 1 14 1 4 0 4 44 9 10 18 42 61 104

nve.no . The statistics were provided the by

Catchment km Catchment

2

121 121 55 211 35 150 395 63 16 210 39 63 9 29 381 120 217 171 379 828 525

°C temp. summer Mean

4.3 3.1 4.1 3.7 4.3 4.2 4.4 6.0 6.4 4.5 5.7 5.2 5.1 4.0 4.1 4.5 3.9 4.5 4.8 4.5 4.7

°C temp. winter Mean

-4.2 -5.1 -4.5 -6.1 -7.9 -6.9 -5.9 -1.8 -1.4 -4.0 -1.6 -2.0 -2.7 -3.3 -4.2 -3.2 -5.3 -5.9 -5.8 -7.8 -6.1

mm Mean summer eposition eposition summer Mean

594 504 434 406 356 356 375 387 447 435 423 460 489 518 590 614 586 442 400 390 380

mm Mean winter deposition deposition winter Mean 1000 832 725 571 497 527 669 809 944 900 919 984 938 1013 1062 1151 984 742 582 571 434 (W – 11 western(W – 11 catchments, central C/E – 16 and eastern catchments) on Hardangervidda ). The). vital statistics the of catchments were calculated using the NEVINA procedure (NVE Atlas, nve.no Limits catchment of Confluence Austdøla of and river from Lake Confluence Skykkjedalselvi/ of Nordelvi Upstream Stakseng Upstream outflow Lake Sysenvatn reservoir Upstream Nybu Upstream Nybu Upstream Valursfossen Upstream Erdalsstølen Upstream Tverrlii Upstream Søtefossen Confluence with Kinso Upstream confluence Opo/ of Skrikjo Upstream Brattespe Upstream outflow Vendevatnet Lake Vetle Upstream outflow Lake reservoir Upstream outflow Lake Reinsnosvatnet Upstream outflow Lake Valldalsvatnet reservoir Upstream outflow Lake Bordalsvatnet reservoir Upstream outflow Lake Songa reservoir Upstream Mogen Tributaries upstream Lake Møsvatn reservoir ecscl. Kvenna Lepidurus arcticus

Catchment name Austdøla Sima Isdølo Leiro Svinto Veig Erdalsvassdr. Bjotveitelvi Kinso Vivippo Opo Espeelvi Vendo Tysso Ljoso Austdølo/ Suldalsvassdr. Bora Songa Kvenna Møsvatn Catchment no. Catchment 1 2 3A 3B 3C 4 5 6A 6B 7A 7B 8 9A 9B 10 11 12 13 14 15 16

The catchments 27 surveyed for Region W W C/E C/E C/E C/E C/E C/E W C/E W W W W W W W W C/E C/E C/E Table 1. Table 1. ResourcesWater and Energy Directorate ( in Appendix 1. 80

Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

L. arcticus L.

with with water chemistry data (97%) are from the period July – October Proportion (%) of lakes lakes of (%) Proportion (Appendix 1). In many lakes, however, only single values of 93 67 83 83 100 100 57 calcium are available. We have obtained water chemistry data

arcticus (concurrent values of calcium concentration and pH) from the L. L. with lakes of No.

13 2 5 43 5 4 136 literature for 237 lakes on Hardangervidda (Appendix 1).

The climatic conditions on Hardangervidda are highly No. of lakes examined lakes of No. variable (Qvenild et al. 2018). We used the NEVINA procedure

14 3 6 52 5 4 238 to compute winter and summer deposition and winter and

summer temperatures as mean values for the normal period No. of lakes of No. 1961 – 1990 (NVE Atlas, nve.no). The winter deposition 156 39 37 154 10 19 930 (October – April) differs substantially between the western and

eastern parts of the plateau, being 1151 mm in Austdølo/Ljoso Lake area km area Lake 2

102 10 22 134 3 16 656 catchment and 292 mm in Uvdalselvi catchment, respectively

(Table 1). Mean summer deposition values (May – September)

Catchment km Catchment are less divergent, being 614 and 332 mm, respectively. 2

732 196 188 1179 61 76 6569 In most Hardangervidda lakes and rivers, brown trout is

the only fish species present. Salvelinus alpinus

°C Mean summer temp. temp. summer Mean (Linnaeus, 1758) and European minnow Phoxinus phoxinus

4.8 5.1 4.8 4.7 5.0 4.6 (Linnaeus, 1758) are coexisting with brown trout in some of the lakes (Appendix 1). Arctic char is found in 3.1% (N=29) of the

th °C Mean winter temp. temp. winter Mean 930 named lakes, mainly due to stockings in the 20 century.

-6.7 -6.6 -6.8 -8.1 -5.8 -7.4 In the Lågen catchment, they were stocked in Lake Breidvatnet,

mm a headwater lake in the tributary Æanåi in 1910 (Dahl 1920), Mean summer eposition eposition summer Mean from where they spread downstream and to some neighbouring

339 332 355 344 323 339 lakes in the following years. A dense population of Arctic char that emerged in Lake Breidvatnet affected the local crustacean

mm community (Dahl 1920). More recently, Arctic char have also Mean winter deposition deposition winter Mean

304 292 314 511 322 404 established themselves in the Leiro catchment (Rognerud et al. 2003). The European minnow has been introduced to 4.5% (N=42) of the 930 lakes since the 1970s. It was first observed in Lake Ørteren reservoir in 1973 (Anonymous 1974), from where it spread to many of the lakes in the north-eastern part of the plateau. Mapping of Lepidurus arcticus Data on the presence or absence of L. arcticus were screened from technical reports of varying standards and from some scientific papers (Appendix 1). like L. arcticus may be difficult to detect using conventional benthic sampling equipment at low densities and/or patchy distribution (Fjellheim et al. 2007). Methods used in sampling L. arcticus on Hardangervidda include the Ekman bottom sampler (Dahl 1917;

Limits catchment of Upstream Lake Gøyst reservoir Upstream confluence Jønndølå Tøddøla/ Upstream outflow Lake Sønstevatn reservoir Upstream outflow Lake Orsjoren Upstream confluence Ufysja/ Ljosevassåi Upstream outflow Lake Ørteren reservoir Amundsen 1976), Petersen sampler (Amundsen 1976), plankton sieves (Halvorsen 1973), artificial substrates (jute bags) and benthic littoral kick samples (Walseng et al. 1994; Walseng et al. 1996; Fjellheim et al. 2007). However, in most publications, the occurrence of L. arcticus is mainly based on brown trout stomach analyses.

Catchment name Gøyst/ Mår Uvdalselvi Ølmøsåi Lågen Ufysja Ørteråni TOTAL Many reports contain only a note referring to a finding of

L. arcticus. In more detailed investigations, the occurrence of Catchment no. Catchment different food items is normally given as volume % based on 17 18A 18B 19A 19B 20

stomach analyses. In this study, the occurrence of L. arcticus

Continued. Region is only noted as presence or absence of the species. In this

C/E C/E C/E C/E C/E C/E context, we assume viable populations of L. arcticus in lakes Table 1. Table 1. 81 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

where the species have been noted at least once. In lakes within the 27 catchments studied (Appendix 1, Table 1). Their with negative findings of L. arcticus they may still occur as pattern of distribution is highly skewed, with almost all the findings are highly sensitive to the method used (Amundsen records coming from the 16 central and eastern catchments 1976; Fjellheim et al. 2007), time of sampling (Aass 1969; (Figure 2). In this area, 775 named lakes were identified. In the Hesthagen 1979; Qvenild et al. 2018), thermal conditions (Aass 181 natural lakes examined, L. arcticus was detected in 69% 1969; Simonsen & Valderhaug 1994; Borgstrøm 2016; Qvenild (N=124). Lepidurus arcticus occurred in all ten reservoirs in et al. 2018), and predation from dense fish populations (Dahl this area. 1920; Aass 1969; Borgstrøm et al. 1985; Qvenild & Rognerud In the 11 western catchments, 155 named lakes were 2018). This is emphasised by results from lakes repeatedly identified. Here, L. arcticus was searched for in 47 lakes, but examined that produced both positive and negative findings recorded in only two (4.0%), both in the Austdøla catchment (Hesthagen 1979; Qvenild & Rognerud 2018). As most of the in the north-western region (Lake Langvatnet and Lake findings of L. arcticus are from stomach analyses, the records Austdalsvatnet). Thus, in the remaining ten catchments in this concern adult specimens. With hauls, different instar stages and region, L. arcticus has never been recorded. nauplius larvae were also detected (Halvorsen 1973; Walseng et al. 1996). The significance of altitude and lake size We analysed the occurrence of L. arcticus in relation to altitude and lake size from natural lakes within the 16 catchments in RESULTS the central and eastern parts of the plateau. The 124 lakes that Distribution of lakes hosting Lepidurus arcticus contain L. arcticus in this area lie at altitudes between 832 and Lepidurus arcticus was detected in 136 of 238 lakes (57%) 1386 m a.s.l., most of them (77%) between 1100 – 1299 m a.s.l.

Figure 2. Lakes surveyed for the presence of Lepidurus arcticus are shown by dots, with positive findings shown by filled dots. Details of the localities are given in Appendix 1. Norwegian Mapping Authority CC BY 4.0.

82 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

(Table 2). From an altitude of 900 m a.s.l., the proportion of L. arcticus was found in 97% of the 62 lakes ≥1.0 km2, while lakes containing L. arcticus rose to its highest level at 1100 – the species was found in only 54% of the 119 lakes <1.0 km2. 1199 m a.s.l., where 80% contained L. arcticus. In total, 50 and The smallest lake with L. arcticus was Lake Nedre Vivevatn 72% of the lakes lower and higher than 1100 m a.s.l. hosted L. (0.023 km2). arcticus, respectively. Lake Kolsnutgryslane at 1386 m a.s.l., in the Kvenna catchment, is the highest known locality hosting L. Occurrence of Lepidurus arcticus in relation to climate arcticus on Hardangervidda. Of the 26 lakes at higher altitudes, conditions and water chemistry only two lakes have been surveyed for L. arcticus, and both The 11 catchments in the western part of the plateau with returned negative findings. Of the 19 lakes located at altitudes scarcely any L. arcticus had high winter depositions of snow, below 1000 m a.s.l., ten were surveyed, and five of them with a mean value of 961±109 mm (Table 1). Lakes in these contained L. arcticus. catchments were consistently low in calcium, with a mean The 765 natural lakes in this area ranged in size from 0.004 concentration of 0.81±0.48 mg L-1 (range 0.32 – 2.62 mg to 18.38 km2, most of them covered an area <1.0 km2 (90%). The L-1, N=36) (Appendix 1). Calcium concentrations in the two probability of occurrence of L. arcticus increased significantly localities with L. arcticus in this area, lakes Langvatnet and with lake size as shown by a logistic regression: p(occurrence) = Austdalsvatnet, were 0.32 and 1.20 mg L-1, respectively. Of the (1+exp (1.380 + 2.026 ∙ log Area))-1 (likelihood-ratio chi-square 36 localities, 72% had calcium concentrations <1.0 mg L-1. The test=31.12, p<0.0001) (Figure 3). Of the 181 lakes examined, concurrent pH in these lakes ranged from 5.35 to 7.00, with 19% lying below 6.0. The TOC values are extremely low, with all Table 2. Frequency (%) of lakes with Lepidurus arcticus at values ≤0.6 mg C L-1. different altitudes in the 16 central and eastern catchments on On the other hand, in the 16 catchments in central and Hardangervidda. eastern areas in which L. arcticus is common, the mean winter

- deposition is 527±183 mm (Table 1). Concurrent values of calcium concentration and pH were available from 201 lakes (Appendix 1), with a mean calcium concentration of 1.61±1.12 mg L-1. The minimum and maximum calcium values for the lakes (%) lakes in this area were measured in Lake Svartavasstjørni Altitude m a.s.l. natural of No. lakes lakes Examined Lakes with Lepidurus arcticus Proportion of Lepidurus arcti cus (Lågen catchment) and Lake Nøkkatjørni (Bjoreio catchment), <900 4 4 3 75 with 0.28 and 5.73 mg L-1, respectively. In this area, 37% of the 900-999 15 6 2 33 lakes had a calcium concentration <1.0 mg L-1. The concurrent 1000-1099 59 18 9 50 pH varied from 5.19 to 7.28, with 10% <6.0. In this area, L. 1100-1199 264 66 53 80 arcticus is recorded in 95 lakes in which the mean calcium 1200-1299 270 57 42 74 concentration was 1.67±1.14 mg L-1 (range 0.28 – 5.54 mg L-1). 1300-1399 141 30 15 50 The concurrent pH values ranged from 5.49 to 7.26 also with ≥1400 12 0 0 0 11% of the values <6.0. Also in this area, the TOC values are -1 765 181 124 69 low, as 85% (N=52) of the localities were below 2.0 mg C L . Winter deposition and calcium concentration were significantly different between the western and the central/ eastern areas (likelihood-ratio chi-square test=120.91 (p<0.0001) and 20.61 (p<0.0001)), respectively (Figure 4). At calcium concentrations <1.0 mg L-1, no positive findings of L. arcticus were found in areas with snow deposits >700 mm. Both winter deposition (Wd) and calcium concentration (Ca) could be used to characterise the western and central/ eastern areas respectively. The probability of being categorised as a lake in the central/eastern area could be modelled by a logistic regression: p = 1 - (1 + exp (15.158 + 4.278 ∙ log Ca - 0.019 ∙ Wd))-1 (likelihood-ratio χ2 test = 169.7, p<0.0001).

Figure 3. Logistic regression curve showing the probability of occurrence of Lepidurus arcticus as a function of lake surface area. Of the 181 lakes studied, Lepidurus arcticus was found in 97% of the 62 lakes ≥1.0 km2, while the species was found in only 54% of 119 lakes <1.0 km2. 83 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

L. arcticus. In a similar study in two lakes, artificial substrate (jute bags) and benthic littoral kick sampling were used in addition to stomach sampling (Fjellheim et al. 2007). In these lakes, where L. arcticus appeared in low numbers, only fish stomach analyses had positive findings. Stomach analyses thus proved to be far the most effective method to detect the presence of L. arcticus, especially when L. arcticus appears in low number and/ or has a patchy distribution. Even though adult L. arcticus is a big food item, it is not necessarily easy to catch for the brown trout, especially not to the small sizes. It was a significant positive correlation between the frequency (%) of L. arcticus in the fish stomachs and fish size (Qvenild et al. 2018). In fishery investigations, multimesh gillnets are frequently used. In clear water mountain lakes, small brown trout tends to pose the dominant part of the catch in such gears. If so, the probability to detect L. arcticus may be lower. In our material, 423 investigations can be properly dated, -1 Figure 4. Scatter plot of calcium concentration (mg L ) and winter 102 of which delivered negative findings in lakes known to deposition (mm) for the lakes in the two catchment areas. Bivariate host L. arcticus. This may be due to timing (early sampling), 90% confidence ellipses are shown for each of the two areas. methods used, low density and/ or patchy distribution of L. arcticus. As 44% (N=102) of these studies took place in June DISCUSSION – July, timing seems to be crucial. Most fish investigations in high mountain lakes where L. arcticus is expected as a food In 1910, the first six localities with L. arcticus in the north- item take this into account. Of the 423 investigations, 80% took eastern part of Hardangervidda were documented (Dahl 1917), place in August to October. and by 1940, only 20 lakes with L. arcticus had been identified Although L. arcticus may thrive at low temperatures, (Dahl 1917; S. Sømme 1934). This came to 50% of all a minimum thermal input is needed for their growth into Norwegian lakes that were known to host L. arcticus at that adult stages (Qvenild et al. 2018). This is emphasized in cold time. In the 1970s, enough investigations had been conducted summers where L. arcticus has a delayed appearance as brown to trace a rough distribution pattern on Hardangervidda trout food (S. Sømme 1934; Aass 1969; Pedersen & Scobie (Anonymous 1974). Lepidurus arcticus is well established in 1990; Simonsen & Valderhaug 1994; Borgstrøm 2016). In cold the southern mountain range in Norway in the alpine zone summers such as those of 2012 and 2015 they may not be seen (zone above the tree line) and to some extent in the subalpine until September, if at all (Borgstrøm 2016; Qvenild et al. 2018). zone (mainly the birch zone), indicating that temperature In an extensive study covering 236 fishing events from five preferences limit its distribution. However, in the western lakes on Hardangervidda in 2003 – 2016, the fishing events part of Hardangervidda it must be additive vectors. This study with positive findings covered a period from 27 July (in 2009) focuses on the environmental conditions, in order to elucidate to 2 October (in 2006). In the surveyed literature (cf. references why L. arcticus is missing in this region. in Appendix 1), 10% of those with positive findings were from July. Only in one investigation was L. arcticus detected The significance of the sampling method earlier than 18 July (11 July in Lake Isdalsvatnet; Lehman In many lakes, different kind of bottom samplers are used & Wiers 2004). This confirms existing experience that L. simultaneously with fish sampling (Dahl 1917; Amundsen 1976; arcticus basically contributes to the brown trout diet in August Fjellheim et al. 2007). In a test-fishing programme that covered – September on Hardangervidda (Dahl 1917; I. Sømme 1941). 15 lakes hosting L. arcticus, bottom samplers (Ekman or/and In the 16 central and eastern catchments, L. arcticus was Petersen) were used simultaneously with test fishing (Amundsen recorded in 70% of the 191 lakes examined (including the ten 1976). In only one lake (7%), L. arcticus was detected by the reservoirs). Lepidurus arcticus was recorded in 55% of the bottom samplers, compared with 12 lakes (80%) from which 124 lakes sampled once or twice. With more than two repeat stomachs were analysed. In a monitoring programme with samplings, almost all gave positive records (99%). This was benthic littoral kick sampling in 18 lakes hosting L. arcticus also emphasized in an extensive study in Lake Sandvatn in in 1978, repeated in 1994, L. arcticus was detected in only Kvenna catchment in 2003 – 2016 (Qvenild & Rognerud 2018), one lake in each of the sampling periods (Walseng et al. 1994, where, L. arcticus contributed significantly as brown trout food 1996). In the 1994 investigation, brown trout were also sampled in August in most years. However, in some years they were with multimesh gillnets in six lakes, five of which provided scarcely ever observed. This was partly due to cold summers

84 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

(2012, 2015) and partly to predation pressure from brown trout 1969). The resting eggs of L. arcticus tolerate both freezing (2006, 2007). Thus, repeated sampling is essential in detecting and drying (Dahl 1932; Longhurst 1955; Aass 1969; Blomkvist L. arcticus. 1995). Thus, they are well adapted to the unstable conditions Fish predation may serious reduce L. arcticus populations. in the littoral zone in reservoirs with variable water levels. On Hardangervidda this has been shown for brown trout However, in this context, temperature conditions seem to be (Qvenild & Rognerud 2018), Arctic char (Dahl 1920; Aass crucial, as was revealed in Lake Pålsbufjorden at 749 m.a.s.l. 1969) and European minnow (Rognerud et al. 2003). When lower in the Lågen catchment (Dahl 1932). The regulation one or more of these species are present, L. arcticus may be regime turned the lake into a colder habitat suitable for the difficult to detect. development of L. arcticus (Aass 1969). The water level amplitude in hydropower reservoirs does The distribution pattern and potential for new observations not seem to be decisive (Aass 1969). All the ten reservoirs in of Lepidurus arcticus the central and eastern parts of Hardangervidda still host L. Hardangervidda marks the southernmost distribution limit of L. arcticus despite water amplitudes up to 66 m (Lake Sysenvatn). arcticus, although a few localities in the neighbouring mountain In the western catchments, no records of L. arcticus after area further south are known (S. Sømme 1934; Borgstrøm et al. impoundment are known. Consequently, there must be other 1976; Økland & Økland 2003; Artsdatabanken.no). This study factors that limit their occurrence in reservoirs in this part of revealed 136 lakes hosting L. arcticus (cf. Table 1). Almost the plateau. all the lakes are located in the central and eastern part of the The presence of a species is a function of the proximity mountain plateau. Our study has thus confirmed the pattern of suitable refugia. In this study, the occurrence of L. arcticus of distribution seen in the beginning of the 1970s (Anonymous increased significantly with lake size, which demonstrates 1974). However, it is now based on far more localities. that this is a factor of great significance, as access to refugia In the 16 catchments at the central and eastern part of the in larger and deeper lakes tends to be better. In this context, plateau where L. arcticus is common, 25% of the named lakes this seems to be a crucial factor, relative to both water quality (N=775) have so far been properly surveyed. This particular and fish predation (Fjellheim et al. 2007; Qvenild & Rognerud crustacean can be difficult to catch, and investigations that 2018). include more than two repeat samplings may be needed to be In addition to the 930 named lakes on Hardangervidda, certain of detecting it. Of the 191 lakes, only 35% have been there are large numbers of unnamed lakes and ponds. In the examined more than twice and the potential for new findings high Arctic, L. arcticus is common in such localities although of L. arcticus is obvious. Hence, the estimate that at least 136 they may freeze solid throughout the winter (Lakka 2013). localities host L. arcticus in this area should easily be increased Lepidurus arcticus have also been recorded in some small by more targeted surveys. lakes and ponds on the Scandinavian peninsula (Berg 1954; The distribution of L. arcticus throughout the Norwegian Hesthagen 1979; Blomkvist 1995; Walseng et al. 1996). Even in mountain range, which decreases to the north, is mainly such localities, L. arcticus seems to be an exception (S. Sømme explained by their temperature preferences (S. Sømme 1934; 1934; Halvorsen 1973; Blomkvist 1995; Klausen 2012; Walseng Aass 1969; Økland & Økland 2003). The general drop in air et al. 1996). In most small lakes and ponds, the environmental temperature, which falls by about 0.6°C per 100 m altitude conditions may be too variable and lack the necessary refugia correlates with a corresponding drop in water temperature to provide survival conditions for L. arcticus. Both temperature (Kvambekk & Melvold 2010; Lindholm et al. 2015; Qvenild and acid water may have detrimental impacts in such localities et al. 2018). Hence, a thermal deficit in the input needed to (Olofsson 1918 cited in Blomkvist 1995; Fjellheim et al. 2001; complete its reproductive cycle may limit the species at high Lakka 2013). Another branchiopod, palludosa altitudes (cf. Aass 1969: Qvenild et al. 2018). This may be O.F.Müller, 1758, has been extinct in many ponds on its reflected in the maximum frequency of L. arcticus found southernmost border in the Dovrefjell mountain area since the in lakes at altitudes of 1100 – 1299 m a.s.l (cf. Table 2). On 1970s, which was related to elevated summer temperatures Hardangervidda the highest lake hosting L. arcticus is Lake (Lindholm et al. 2012, 2015). Lepidurus arcticus may also be Kolsnutgryslane at 1386 m a.s.l. However, they may exist at susceptible to rises in temperatures in the upper part of their higher altitudes as the species has been recorded up to 1518 m thermal range (Arnold 1966; Klausen 2012; Lakka 2013). a.s.l. nationwide (Økland & Økland 2003). On Hardangervidda only 26 lakes (3.4%) are sited at higher altitudes than 1386 m Environmental constraints on the distribution of Lepidurus a.s.l. and only two of them have so far been examined. arcticus Lepidurus arcticus normally reach their maximum On Hardangervidda, the western area is extremely barren abundance in the alpine zone in Southern Norway (S. Sømme with only a thin layer of moraine or glacio-fluvial deposits on 1934; Aass 1969; Økland & Økland 2003). However, their Precambrian bedrock, almost without vegetation and with a distribution may extend into the subalpine zone, especially in bedrock of granitic and dioritic gneisses, giving dilute water lakes regulated for hydropower production (Dahl 1932; Aass low in calcium (0.81±0.48 mg L-1). The catchments in the

85 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

central and eastern areas where L. arcticus commonly occurs, into adults (Qvenild et al. 2018). High winter deposition, late has a thicker moraine cover with more vegetation, and the water ice break-up and low water temperatures in a short growing quality is much better with respect to calcium (1.61±1.12 mg season may seriously impede the growth and development of L-1). In the western and central/ eastern part, 72% and 38% of L. arcticus. Evidence for this was documented for a number of the lakes have calcium concentrations <1.0 mg L-1, respectively. individual years on Hardangervidda (Pedersen & Scobie 1990: The lower lethal threshold for calcium demand has yet Simonsen & Valderhaug 1994; Borgstrøm 2016; Qvenild et al. to be established for L. arcticus. The species is also highly 2018). Thus, the combination of low thermal input and dilute sensitive to acid water, becoming extinct in waters with waters in this area may result in excessively hostile conditions pH below 5.5 (Borgstrøm & Hendrey 1976; Fjellheim & to L. arcticus. When calcium concentrations are <1.0 mg L-1, Raddum 1990; Lakka 2013). Rapid postmoult calcification of L. arcticus was not found in areas with snow deposits >700 the exoskeleton is essential to all crustaceans, and low calcium mm. (cf. Figure 4). However, L. arcticus may maintain viable concentrations may impede ecdysis (Rukke 2002). This effect populations even in some lakes in areas with snow deposition may be reinforced by low pH levels, which often interfere >700 mm. Lake Busetevatn in the Erdalsvassdraget catchment with low calcium concentrations. Almost all the 29 lakes and lakes in the Kinso catchment host L. arcticus despite winter with pH <6.0, ten of which hosted L. arcticus (cf. Appendix depositions of ~900 mm. However, these lakes have relatively 1) had calcium concentrations <1.0 mg L-1. In these lakes, L. high calcium concentrations (>1.4 mg L-1). Hence, L. arcticus arcticus may cope with such extremes and must be regarded is able to complete its life cycle even with snow deposits >700 as “living on the edge” of survival. At such critical levels, mm, if calcium levels are sufficiently high. access to proximity refugia is assumed to be crucial. In Lake Svartevatnet in Lågen catchment, L. arcticus was not recorded Climate change – what will be the consequences for Lepidurus until the water quality was improved by a liming programme arcticus ? that started in 1994 (Fjellheim et al. 2001). This was assumed For the Norwegian mainland as a whole, there has been a not to be an effect of the stocking of L. arcticus, but rather general increase in precipitation since the 1980s (Hanssen- of access to refugia within the lake with better water quality. Bauer et al. 2017). This pattern has also been observed on Hence, the lower thresholds to pH and calcium should exceed Hardangervidda, where the annual winter deposition has the measured level in Lake Svartevatnet in 1993, i.e. prior to increased, including an increase in the number of snow-rich liming. In that year, the summer calcium concentration and winters with a decrease in the difference from west to east pH were 0.35 mg L-1 and 5.69, respectively. This indicates that (Qvenild et al. 2018). In addition to later ice break-up and colder ambient calcium concentration levels ~0.5 mg L-1 is near to the waters, elevated precipitation levels will result in higher run-off lower lethal threshold for L. arcticus. and more dilute waters (Presthus Heggen et al. 2010. This study has clearly shown that L. arcticus can thrive in A similar significant increase in summer air temperatures waters lower in calcium than <1.0 mg L-1 and also with pH <6.0. since the 1980s has also been documented on Hardangervidda Even though the lakes in the two areas have quite different levels (Qvenild et al. 2018), and a further increase is projected toward of calcium, many of them with positive findings of L. arcticus the end of the century. As water temperature is closely linked to in the central/ eastern area lie within the lower range of the air temperature this will also affect the aquatic biota (Kvambekk scale, with 35 lakes having calcium concentrations <1.0 mg L-1, & Melvold 2010; Lindholm et al. 2015; Qvenild et al. 2018). In and seven of them <0.5 mg L-1. Thus, the calcium concentration shallow lakes and ponds, the water temperature may exceed the in most of the lakes in the western area should not be critical upper tolerance limit of L. arcticus, which is estimated to be to L. arcticus survival and in this part of Hardangervidda, 19 – 20 °C (Arnold 1966; Klausen 2012; Lakka 2013), although additional vectors determine their absence or presence. it may be higher for short duration exposures. In deeper lakes, There is a steep gradient in snow deposition from west to the water temperature seldom passes these tolerance limits for east, with 1151 mm in the Austdøla/Ljoso catchment in the more than a few days, and only in the epilimnion (Qvenild et west, compared to 292 mm in the Uvdalselvi catchment in al. 2018). Hence, lake size would appear to be important for L. the east (cf. Table 1). All the western catchments have snow arcticus survival in a future warmer climate. deposits >700 mm. Heavy snow deposits seriously affect the The ecological sensitivity is high in mountain areas, where thermal conditions in the lakes (Borgstrøm 2016; Qvenild et many species are living at the thresholds of their tolerance and al. 2018). The large amounts of snow deposited in western close to their geographical limits. Hence, protracted warm areas imply that the ice break-up usually takes place much later periods may be problematic for a cold-adapted species like L. than in eastern areas (Borgstrøm 2016; Qvenild et al. 2018). arcticus at the southernmost outreach of its distribution range. Hatching of L. arcticus normally coincides with snow melt Even though L. arcticus may tolerate high temperatures for and ice break-up (Halvorsen 1973; Simonsen & Valderhaug some time, the accumulated thermal input may exceed the 1994; Fjellheim et al. 2007; Borgstrøm 2016). Though low upper thermal threshold and result in mismatches between temperature is not a problem to an arctic species like L. environmental conditions and life-cycle events. This may force arcticus, a minimum thermal input is needed for development range retractions of L. arcticus from the lower part of their

86 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019 distribution area, as has already been experienced by another on trout in Lake Paalsbufjord. Skrifter utgitt av Det Norske crustacean species, B. paludosa (Lindholm et al. 2012, 2015). Videnskaps-Akademi i . I. Mat.-Naturv. Klasse. 1931. No. From a conservation point of view, detailed mapping of the 4. 53 pp. present distribution of L. arcticus on Hardangervidda should be Fjellheim A, Raddum GG. 1990. Acid precipitation: biological monitoring of streams and lakes. The Science of the Total highly rewarding. Environment, 96: 57–66. Fjellheim A, Tysse Å, Bjerknes W. 2001. Reappearance of highly acid sensitive invertebrates after liming of an alpine lake ACKNOWLEDGEMENTS ecosystem. Water, Air and Soil Pollution 130: 1391–1396. Fjellheim A, Tysse Å, Bjerknes V, Wright RF. 2002. Finprikkauren The authors are grateful to Eirik Fjeld for valuable help with the på Hardangervidda. DN-utredning 2002-1. 58 pp. (In statistics, and Roar Kiær for drawing the maps. Norwegian). Fjellheim A, Tysse Å, Bjerknes W. 2007. Fish Stomachs as a Biomonitoring Tool in Studies of Invertebrate Recovery. Water, Air and Soil Pollution 7: 293–300. doi: 10.1007/s11267-006- REFERENCES 9074-x Aass P. 1969. Crustacea, especially Lepidurus arcticus Pallas, as Fjellheim A, Tysse Å, Gåsdal O, Stakseng H. 2018. Finprikkauren brown trout food in Norwegian mountain reservoirs. Institute på Hardangervidda. UNI Research Miljø – LFI, Univ. i Bergen. of Freshwater Research, Drottningholm. Report No 49, 1969: Årsrapport 2017. 14 pp. (In Norwegian). 183–201. Hanssen-Bauer I, Førland EJ, Haddeland I, Hisdal H, Lawrence D, Amundsen T. 1976. Fiskeribiologiske undersøkelser for Mayer S, Nesje A, Nilsen JEØ, Sandven S, Sandø AB, Sorteberg Dagalivassdraget 1974–75. Fiskerikonsulenten for Øst-Norge. A, Ådlandsvik B. 2017. 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88 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 55, 124 55, 124 124 86, 51, 124 124 124 51 51, 55 51, 55 55 55, 124 55, 124 51 51 55 124 124 56, 71 67 56, 1, 124 67, 56, 120, 121 not conf. not conf. not conf. not conf.

TOC mg C L C mg TOC

-1

Ca mg L mg Ca

-1

0.32 0.40 0.64 0.80 1.68 1.20 0.52 0.32 1.16 1.00 1.48 2.12 0.88 0.82 pH

6.90 6.00 5.60 6.10 7.00 6.90 6.80 6.90 6.90 6.70 6.65 6.52 6.60 6.02 Date

06.08.71 09.09.68 07.09.68 11.09.68 09.08.71 08.08.71 07.08.71 08.08.71 27.07.67 16.08.67 13.09.08 14.06.88 29.07.67 01.06.98 No. pos. rec. pos. No.

0 0 2 0 0 1 0 0 0 0 0 1 2 0 No. exam. No. (W – 11 western(W – 11 catchments, central C/E – 16 and eastern 2 2 3 1 0 0 0 1 1 1 2 2 0 0 1 0 0 2 2 3 1 0 0 0 0 6742742 6744428 6739016 6745937 6747708 6745100 6743690 6743724 6742930 6743018 6743002 6742080 6741101 6740013 6740315 6735115 6740581 6731638 6726176 6722461 6728945 6729452 6729200 6729101 6728526 EPSG:25833 Lepidurus arcticus

70590 72020 69287 74117 73417 70313 80149 72907 75886 69545 68971 68552 70461 72883 68354 76463 76822 75584 76533 82835 95280 90932 92115 91807 91182

P. phoxinus P. S. alpinus S.

x x x x x x

Max. depth m depth Max.

Min. depth m depth Min.

Area km Area

2

0.63 1.28 6.40 0.53 1.05 0.19 0.16 0.08 0.32 0.09 0.06 0.13 0.35 0.13 0.12 1.29 0.45 0.43 1.07 10.42 1.23 0.15 0.06 0.10 0.26

m a.s.l. m Altitude Altitude

1040 1040 1158 1140 1153 1081 1263 1059 1163 954 936 907 1194 1192 1107 905 1216 837 832 880 1190 1227 1165 1165 1154 Reg. x x x x x Catchment Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Austdøla Sima Sima Sima Isdølo Leiro Leiro Leiro Leiro Leiro Leiro

Lake Austdølnutvatnet Rundavatnet Langvatnet Kvilinganutvatn Grøndalsvatnet S. Floskefonnvatnet Memorgevatn V. Austdalsvatnet Austdalsvatnet Rossevatni Rossevatni Austdølvatnet Olbogavatnet Reipstjørnane Grasbotntjørni Rembesdalsvatn Holmavatn Skykkjedalsvatn Isdalsvatnet Sysenvatn Finsbergvatnet Lassheldretjørni Lassheldretjørni Lassheldretjørni Lassheldretjørni NVE ID NVE indicates that a lake is regulated. The number examinations of is specified together with any associated positive records. The numbered references are the given below table. 1917 1918 1921 1922 1923 16623 16637 16656 16664 16677 16682 16712 16747 16781 16787 1914 1915 17050 1906 1907 17070 17084 17092 17095 17114

Reg Catchm. no. Catchm.

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 2 3A 3B 3B 3B 3B 3B 3B

. Specific information lakes about in 367 catchments, 27 which were of surveyed 238 the for occurrence of Region W W W W W W W W W W W W W W W W W W C/E C/E C/E C/E C/E C/E C/E Appendix 1 catchments). 89 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 122121, 79 25, 79 79 25 25 25 25, 79 42, 86 86, 83, 79, 100 100 79,83,86 79, 67, 27, 100, 86, 83, 119 79, 100 129 79, 124, 129 124 129 100, 79, 79 10, 79, 100 79, 128 79, 128 79, 128 128 100, 79,

TOC mg C L C mg TOC

-1

1,2 1,5 1,2 <0,2 1,0 1,2 0,4

Ca mg L mg Ca

-1

0.62 0.85 1.91 0.92 4.07 0.75 4.85 5.73 4.16 1.68 0.96 1.00 1.67 3.45 3.32 pH

6.03 6.38 6.51 6.46 7.00 5.93 7.10 7.26 7.05 7.00 7.03 7.23 7.26 Date

05.08.99 04.08.97 04.08.97 04.08.97 17.08.04 04.08.97 09.10.97 09.10.97 12.10.97 09.10.97 27.07.67 27.07.67 09.10.97 09.10.97 09.10.97 No. pos. rec. pos. No.

0 1 0 1 1 1 1 3 4 6 0 0 2 0 0 0 2 0 1 1 2 No. exam. No. 1 1 1 2 0 1 0 2 2 4 0 4 6 1 1 3 1 2 1 3 2 2 2 2 6726281 6722876 6723206 6721531 6721561 6721458 6721189 6719647 6706801 6703503 6710029 6701871 6704204 6716740 6714260 6713510 6713570 6712245 6710060 6709369 6709950 6709412 6709418 6704861 EPSG:25833

91211 96421 92974 95206 91140 90234 90854 92219 94317 86436 80895 89929 90972 58355 61500 62549 64265 58651 61616 62617 56589 59177 57334 59771

P. phoxinus P. S. alpinus S.

x Max. depth m depth Max.

21,0 26,4 30,0 11,0 33,0 14,5 34,0 Min. depth m depth Min.

6,7 5,5 7,7 5,9 12,3 5,5 8,6

Area km Area

2

0.74 0.16 0.12 0.64 0.29 0.14 0.12 0.25 0.05 2.65 0.12 0.82 4.54 0.42 0.26 0.85 0.16 0.44 0.11 0.64 0.07 0.50 0.22 1.65

m a.s.l. m Altitude Altitude

1124 1259 1153 1175 1125 1141 1126 1173 1220 1222 1353 1215 1213 1222 1082 1041 1009 1195 1060 1045 1196 1152 1193 1231 Reg. Catchment Leiro Leiro Leiro Leiro Leiro Leiro Leiro Leiro Svinto Bjoreio Bjoreio Bjoreio Bjoreio Veig Veig Veig Veig Veig Veig Veig Veig Veig Veig Veig

Lake Halnetjørni Dalboretjørni OlavsbuvatniI. Skardstjørnane Svartetjørni Skardstjørnane Dyratjørnane Stigstutjørni Langavatnet Nøkkatjørni Kleivshovdtjørna Tinnhølen Vassdalsvatni Reinavatnet Vasslivatn Vivetjørnane Langavatnet Nautavad Vatnalivatnet Hanasteinsvatnet Skinhovda Tresnutvatnet Bersavikvatnet NVE ID NVE

17149 17289 17291 17305 17322 17369 17370 17402 17720 1910 17673 17865 27430 17531 17614 17616 17620 17668 17710 17715 17730 17735 17737 17849 Catchm. no. Catchm.

3B 3B 3B 3B 3B 3B 3B 3B 3C 4 4 4 4 5 5 5 5 5 5 5 5 5 5 5

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1 90 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 19, 79 86 79, 25, 19, 100 100 100 100 86, 79, 79 79 48, 53, 15, 82 100 48, 79, 124 79, 124, 129 129 79, 129 79, 124, 128 79, 124 79 79, 100 100 79, 124 79, 124 79, 100 79 79 79, 128

TOC mg C L C mg TOC

-1

0,4 0,2 0,5 0,4 0,3 0,3 0,5 0,4 1,2

Ca mg L mg Ca

-1

2.20 2.96 2.27 1.08 5.54 3.08 1.40 3.16 2.32 3.32 2.32 1.40 2.24 2.12 1.59 3.89 pH

6.50 7.28 6.73 6.53 7.11 7.15 6.90 7.16 6.60 6.79 7.22 Date

17.07.96 09.10.97 09.10.97 28.09.89 09.10.97 09.10.97 30.07.67 05.08.67 07.08.67 10.08.67 09.10.97 31.10.95 26.08.67 26.08.67 09.10.97 09.10.97 No. pos. rec. pos. No.

0 2 1 1 0 2 1 0 1 1 0 2 1 1 0 0 2 1 0 0 0 No. exam. No. 2 3 0 0 0 3 1 1 4 2 2 3 1 2 4 2 2 1 0 2 3 2 2 2 3 6700837 6699894 6701176 6698738 6697312 6695197 6695987 6695491 6693034 6692865 6713675 6726003 6725553 6726935 6706672 6701409 6704912 6697198 6715304 6715537 6715667 6713136 6712693 6712409 6710520 EPSG:25833

75911 76015 59097 65266 62964 73027 71445 61722 63593 61784 64814 58125 57341 52863 53384 55086 56181 59963 54314 48598 49566 54075 49818 50687 54407

P. phoxinus P.

S. alpinus S. Max. depth m depth Max.

22,5 27,0 53,4 23,8 13,3 Min. depth m depth Min.

6,7 7,8 16,0 5,4 4,4

Area km Area

2

0.12 0.20 0.26 0.32 0.28 0.46 0.14 0.34 0.56 0.22 0.02 0.43 0.06 0.44 4.68 2.39 0.26 0.87 0.23 0.30 0.69 0.52 0.21 0.33 0.19

m a.s.l. m Altitude Altitude

1244 1244 1320 1347 1371 1367 1317 1396 1268 1281 1009 884 884 869 1172 1199 1184 1363 1280 889 900 1125 940 955 1170 Reg. Catchment Veig Veig Veig Veig Veig Veig Veig Veig Veig Veig Veig Erdalsvassdr. Erdalsvassdr. Bjotveitelvi Kinso Kinso Kinso Kinso Kinso Kinso Kinso Kinso Kinso Kinso Kinso

Lake Vierslatjørn Heisantjørnane Øvre Solvatnet Langgrøvatnet Meinsvatnet Herrevatnet Nykktjørnane Grytevatnet Grøndalsvatni Grøndalsvatni Nedre Vivevatn Busetevatnet Vetlevatnet Vatnosetvatnet Veivatnet OmkjelsvatnetN. Kinsevatnet Sandvatnet Trollavatnet Rjuvavatnet Stavalivatnet Lonavatnet Kinsevatnet Fodnastølsvatnet Austmannavatnet NVE ID NVE

17975 17997 18002 18103 18157 18224 18228 18287 18409 18434 67665 17300 27433 27428 1912 1913 17878 18214 27475 27476 27478 27495 27500 27510 27523 Catchm. no. Catchm.

5 5 5 5 5 5 5 5 5 5 5 6A 6A 6B 7A 7A 7A 7A 7A 7A 7A 7A 7A 7A 7A

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E W C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1 91 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 79 79, 128 79 124 82, 100 79, 79 79 79 100, 124 79 100 79 79, 100 85 67, 69 91, 67, 18, 125 85, 69, 67, 125 125 100, 85, 85,67, 125 85, 69, 67, 125 85,67, 125 68, 69, 67, 85, 125

TOC mg C L C mg TOC

-1

0,2 0,6 0,5 0,4 0,2

Ca mg L mg Ca

-1

2.44 1.93 0.94 0.71 0.93 1.28 0.64 0.95 0.64 1.03 pH

7.07 6.65 6.14 6.59 6.80 6.38 6.67 6.40 6.67 Date

10.08.67 09.10.97 02.10.97 02.10.97 02.10.97 01.08.10 01.08.10 09.10.97 01.08.10 01.08.10 No. pos. rec. pos. No.

0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 No. exam. No. 1 3 1 1 2 1 1 3 1 1 0 1 1 1 2 6 4 2 3 4 3 5 6710198 6709363 6709620 6705279 6699113 6724099 6723050 6721377 6712854 6716168 6716792 6713653 6709676 6702750 6704733 6694395 6698163 6693281 6699937 6690183 6690302 6690703 EPSG:25833

50169 55661 51466 55531 53915 52528 48533 48434 44930 44724 44306 42663 43328 39059 41667 39127 49307 54087 40547 46031 50202 54458

P. phoxinus P.

S. alpinus S. Max. depth m depth Max.

15,0 36,0 Min. depth m depth Min.

3,4 8,8

Area km Area

2

0.12 0.56 0.19 0.14 0.55 0.66 0.43 0.24 1.25 0.28 0.05 0.19 0.75 0.25 3.64 7.25 2.92 1.39 3.37 2.06 0.68 5.46

m a.s.l. m Altitude Altitude

1012 1186 1057 1179 1202 1183 1122 700 1014 1108 1109 1016 1105 1238 1268 464 1333 1284 1106 1191 1191 1264 Reg. x x x x x x Catchment Kinso Kinso Kinso Kinso Kinso Vivippo Vivippo Vivippo Opo Opo Opo Opo Opo Espeelvi Vendo Tysso Tysso Tysso Tysso Tysso Tysso Tysso Bersåvatnet Nybuvatnet Omkjelsvatnet

Lake Gravdalsvatnet Nasavatnet Sperrådalsvatn Ø. Birgesdalsvatnet Vassdalsvatnet Grytingsvatn Opesjovatnet Kjølevatni Kjølevatni Skriksetvatn Yskjebotnvatnet Mostjørn St. Vendevatnet Ringedalsvatnet Øvre Tyssevatn Nonskardvatnet Øvre Nibbehølen Øvre HåvardsvatnN. NVE ID NVE

27527 27533 27534 27573 27637 27436 27444 27450 1904 27472 27473 27504 27528 27611 1903 1889 1890 1892 1893 1894 1897 1898 Catchm. no. Catchm.

7A 7A 7A 7A 7A 7B 7B 7B 8 8 8 8 8 9A 9B 10 10 10 10 10 10 10

. Continued. Region C/E C/E C/E C/E C/E W W W W W W W W W W W W W W W W W Appendix 1 92 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 85,67, 125 68, 69, 67, 85, 125 85,67, 125 100 100 100 85,67, 125 85 85 85, 69, 67, 125 100 85 67, 85 85 67 85 85 100 79 100 79 79 100 100

TOC mg C L C mg TOC

-1

0,3 <0,2 <0,2 0,2 <0,2 <0,2 0,3 <0,2

Ca mg L mg Ca

-1

0.53 0.59 0.41 0.43 0.64 0.53 0.69 0.38 0.37 0.33 pH

6.28 6.23 6.31 6.29 6.36 6.51 6.08 5.35 5.95 5.66 Date

01.08.10 15.09.97 09.10.97 09.10.97 01.08.10 09.10.97 09.10.97 21.09.95 02.10.97 09.10.97 No. pos. rec. pos. No.

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 No. exam. No. 3 5 3 0 0 0 3 1 4 4 0 1 2 1 1 1 1 0 1 0 1 1 0 0 6698844 6687621 6685761 6688253 6684844 6701774 6697391 6697205 6697460 6696516 6695930 6696107 6695812 6694926 6695188 6694481 6683669 6683280 6675691 6676790 6678046 6676488 6675600 6681121 EPSG:25833

38299 41622 47989 56089 56982 50388 48130 42541 41824 45895 52128 47503 44070 49095 43332 45595 47553 49829 36722 44738 38106 41372 42015 44889

P. phoxinus P.

S. alpinus S.

Max. depth m depth Max.

Min. depth m depth Min.

Area km Area

2

0.88 6.36 3.35 0.52 0.38 1.28 0.43 0.35 0.04 0.84 0.56 0.15 0.15 0.22 0.11 0.18 0.92 0.16 3.35 1.78 0.57 0.26 0.23 0.62

m a.s.l. m Altitude Altitude

1029 1190 1232 1431 1430 1362 1317 1264 1349 1271 1323 1312 1213 1333 1162 1359 1287 1309 594 1223 630 813 978 1219 Reg. x x x Catchment Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Tysso Austdølo Austdølo Austdølo Austdølo Austdølo Austdølo

. Veidedalsvatn

Lake BersåvatnetN. Langevatnet Breidavatn Jukletjørn Juklevatni Illakleivløkene Nedre Tyssevatn Hadletgrøna Hadletgrøna Holmevatn Selsløken VeidedalsvatnN. Stednesvatnet Ø Tyssehølen Reinakolltjørn Hattasteinsvatnet Løkene Reinsnosvatnet Isvatnet Ljosevatn Svartavatn Raudnosvatnet Søre Blåvatnet NVE ID NVE

1900 1901 1902 18713 18887 27600 27650 27661 27662 27663 27668 27669 27672 27677 27680 27682 27758 27762 1702 1703 23143 23161 23168 27797 Catchm. no. Catchm.

10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 11 11 11 11 11 11

. Continued. Region W W W W W W W W W W W W W W W W W W W W W W W W Appendix 1 93 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 70 69, 67, 100 100 80, 77, 50, 105 100 50, 106 50 76, 2, 50, 20, 96,86, 104, 112 107, 50 50, 79 50, 100 50 50 50, 76 50, 100 101 100 50 100

TOC mg C L C mg TOC

-1

0,4 0,2 0,2 0,3 0,5 1,2 <0,2

Ca mg L mg Ca

-1

0.63 0.79 0.60 2.62 0.36 1.08 1.68 1.63 0.76 3.11 2.04 0.62 0.52 0.62 0.53 0.96 2.10 pH

6.33 6.17 5.93 6.50 5.84 5.90 6.50 6.40 6.20 6.60 6.75 6.00 6.22 6.03 6.30 5.80 6.82 Date

01.08.11 02.10.97 02.10.97 15.09.83 21.09.95 15.09.83 15.09.83 12.09.84 12.09.04 15.09.83 02.10.97 12.09.84 02.10.97 13.09.83 02.10.97 15.09.83 02.10.97 No. pos. rec. pos. No.

0 0 0 7 0 1 0 0 0 0 1 0 No. exam. No. 2 0 0 3 0 1 0 13 0 1 1 1 1 2 1 0 1 1 0 6672273 6675590 6676616 6656673 6667242 6665138 6665519 6659180 6664387 6674883 6674859 6673951 6671441 6669624 6667306 6667223 6666718 6665987 6678455 EPSG:25833

50628 52601 50924 70888 63572 68743 66134 82524 72762 72331 67627 74737 70429 81745 81376 85697 87497 81943 64777 P. phoxinus P.

x x

S. alpinus S.

Max. depth m depth Max.

Min. depth m depth Min.

Area km Area

2

7.33 0.48 0.11 7.69 1.89 1.13 0.39 30.01 0.56 1.00 2.07 0.12 0.37 0.42 0.23 0.19 0.88 0.13 0.41

m a.s.l. m Altitude Altitude

745 930 1006 891 1261 1180 1203 974 1108 1255 1161 1348 1149 1307 1289 1285 1195 1242 1401 Reg. x x x x x Catchment Suldals- vassdr Suldals- vassdr Suldals- vassdr Bora Bora Bora Bora Songa Songa Songa Songa Songa Songa Songa Songa Songa Songa Songa Songa Berutjørni

Lake Valldalsvatnet Vivassvatnet Grønhellervatn Bordalsvatnet Holmasjøen Åremotvatni PoddevatnetN. Songa Storhellervatnet Store Urevatnet Hedlevatnet Bamsetjørn Filleherbergvatn Øvre Midtre Berutjørni Gjevdøltjørni Ugletjørni Nedre Berutjørni Tangasjøen NVE ID NVE

1866 11783 23154 55 85 12118 12157 10 25 11734 11743 11786 11904 11933 12024 12029 12030 12092 19129 Catchm. no. Catchm.

12 12 12 13 13 13 13 14 14 14 14 14 14 14 14 14 14 14 14

. Continued. Region W W W W W W W C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1

94 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 60, 88, 59, 101, 126 60, 88 58, 50, 60, 88, 126, 127 100, 126, 127 62 60,50, 90, 92, 93 62,126 82,62, 86, 126, 127 61 90, 127 61, 86, 126, 127 64, 50, 86, 126 82, 86, 79, 126, 127 48, 62, 17, 82, 99, 86, 126, 127 86, 100 100 58, 100 50

TOC mg C L C mg TOC

-1

0,6 1,4 2,1 0,5

Ca mg L mg Ca

-1

1.33 3.74 1.59 2.13 1.46 1.15 2.73 1.43 0.74 0.44 0.59 1.42 1.19 0.66 1.37 0.82 pH

6.63 6.60 6.62 6.78 6.53 6.40 6.70 6.62 6.37 6.00 5.90 6.36 6.56 6.44 6.67 6.22 Date

13.09.83 15.09.83 15.08.95 02.10.97 15.08.95 12.09.84 03.07.78 15.08.95 16.08.95 16.08.95 12.09.84 16.08.95 15.08.95 21.09.95 02.10.97 02.10.97 No. pos. rec. pos. No.

2 3 4 1 4 13 1 4 2 6 2 2 3 5 1 1 0 No. exam. No. 5 4 5 2 5 15 2 4 2 7 3 4 4 8 1 0 1 1 6676114 6677403 6678032 6679965 6679440 6681359 6684108 6684631 6682778 6684863 6669198 6667249 6688631 6686656 6681191 6674617 6673078 6672489 EPSG:25833

94853 89550 85362 75023 71335 82122 77610 66324 93901 90148 93521 93738 74012 63673 101369 82473 89964 83274

P. phoxinus P.

S. alpinus S. Max. depth m depth Max.

11,5 22,1 7,2 13,0 26,0 25,0 Min. depth m depth Min.

3,3 8,2 1,6 2,6 8,1 10,2

Area km Area

2

1.66 2.62 1.29 2.13 2.92 1.57 1.16 5.00 0.97 2.31 1.55 1.40 1.84 1.52 5.11 0.29 0.47 0.46

m a.s.l. m Altitude Altitude

1030 1068 1071 1136 1147 1112 1141 1166 1149 1195 1329 1384 1319 1170 1210 1250 1163 1291 Reg. Catchment Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna

. Bjørnavatnet

Lake Vollevatnet Briskevatnet Gunleiksbuvatnet BjørnavatnetN. Ø Sandvatn Nedre Krokavatn Kvennsjøen Skardvatnet Fjellsjåen Urdevatnet Store Meinsvatn Valgardsvatni Litlosvatnet Gjuvsjåen Brutjørnane Møruvatnet Merakktjørni NVE ID NVE

12 13 14 15 16 17 18 19 38 39 40 41 42 43 78 11729 11763 11808 Catchm. no. Catchm.

15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1

95 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 86, 126, 127 100 86,50, 126 50, 64, 101 50, 64, 101 50, 64, 101 126 25, 79 117 63 79 82, 8, 63, 54, 102, 126 48, 100 11, 48,49,63,126 117 48, 49, 17, 16, 82, 86, 63, 87, 100, 102 100 95, 100 62 48,63,100 90, 98, 61, 126, 127 63

TOC mg C L C mg TOC

-1

0,6 0,4 0,7 <0,2 1,1 0,4

Ca mg L mg Ca

-1

0.46 0.61 0.79 0.69 0.64 3.46 2.40 1.94 2.57 2.33 5.04 0.38 1.50 0.65 2.30 1.25 0.67 pH

6.05 5.92 5.50 5.81 5.78 6.60 6.96 6.76 6.80 6.60 6.70 6.16 6.70 6.31 6.72 6.26 6.30 Date

16.08.95 24.09.86 15.09.83 13.09.83 13.09.83 10.07.78 31.07.03 21.09.94 06.07.78 11.09.97 08.07.78 21.09.94 11.09.97 15.09.97 09.10.97 15.09.97 17.08.95 No. pos. rec. pos. No.

3 0 2 0 0 0 0 0 0 0 2 0 2 9 1 1 0 4 0 No. exam. No. 4 1 3 1 2 2 1 2 0 1 1 8 2 4 0 12 0 1 1 2 7 1 6670387 6669553 6665449 6663497 6663200 6661411 6690732 6689329 6688551 6691313 6689771 6691012 6689682 6690187 6685788 6688435 6688256 6685129 6684881 6686400 6683553 6685608 EPSG:25833

96007 84810 97056 94405 96143 96393 71055 79782 95298 58269 72971 60847 66343 64063 101243 61644 58549 83830 72085 61466 87775 59870

P. phoxinus P.

S. alpinus S. Max. depth m depth Max.

16,0 15,0 13,0 31,0 15,0 Min. depth m depth Min.

8,6 5,7 6,5 6,1 4,7

Area km Area

2

0.57 0.73 0.50 0.52 0.28 0.93 0.44 0.63 0.08 0.43 0.80 0.42 0.81 0.47 0.15 0.61 0.59 0.31 1.21 0.45 0.64 0.74

m a.s.l. m Altitude Altitude

1353 1273 1386 1383 1383 1390 1322 1327 1233 1288 1322 1236 1269 1249 1270 1182 1332 1310 1150 1282 1205 1299 Reg. Catchment Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna

Lake Meinsvatnet V. Ormetjørni Kolsnutgrysline Hokkebrotvatnet Kostveitvatnet Kostveitvatnet Grottjørnanene Reinavatnet Elsjåen Sledalsvatnet Valgardsvatni Krokavatnet Ambjørsvatnet Skavatn Storhellertjørni Kollsvatnet Kollstjørn Blånuttjørnane Krokavatni Vassdalsvatni Dargesjåen Vassdalsvatni NVE ID NVE

11836 11937 12053 12135 12153 12173 18495 18511 18523 18545 18556 18558 18581 18597 18660 18700 18735 18770 18773 18782 18827 18831 Catchm. no. Catchm.

15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15 15

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1

96 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 90, 126, 61, 127 100 50 82 82 Artsdata- banken.no 59, 88 100 62 2, 50 50 50 101 101 50 64 101 9 9 Artsdata- banken.no 50 50

TOC mg C L C mg TOC

-1

1,5 0,9

Ca mg L mg Ca

-1

0.64 1.01 2.04 0.73 0.84 1.31 0.48 0.71 0.78 1.19 0.70 pH

6.31 6.64 6.30 6.39 5.90 6.80 5.40 6.05 5.19 6.20 5.67 Date

17.08.95 02.10.97 15.09.83 02.10.97 15.09.83 12.09.84 12.09.84 13.09.83 13.09.83 15.09.83 13.09.83 No. pos. rec. pos. No.

4 0 1 1 1 1 1 1 0 0 0 0 0 No. exam. No. 5 0 0 1 1 1 2 0 1 2 1 0 0 0 0 1 0 1 1 0 1 1 6682690 6680696 6681005 6682678 6682727 6682042 6676917 6674151 6684438 6655587 6662765 6668293 6664058 6665028 6662626 6663772 6661056 6658142 6658117 6657991 6652780 6653003 EPSG:25833

89249 95370 90674 69955 68807 58483 93458 95786 74249 114017 126347 106073 117722 107009 121236 98755 116638 140859 141341 121187 123124 123664 P. phoxinus P.

x x x S. alpinus S.

x Max. depth m depth Max.

14,0 Min. depth m depth Min.

4,5

Area km Area

2

0.72 0.70 0.35 0.36 0.26 1.75 0.38 0.20 0.93 78.77 6.99 0.86 0.25 0.31 0.42 0.61 0.63 0.22 0.06 0.08 0.24 0.17

m a.s.l. m Altitude Altitude

1255 1148 1205 1282 1288 1204 1045 1303 1141 919 1145 1241 1240 1394 1208 1352 1237 1077 1072 1003 924 923 Reg. x Catchment Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Kvenna Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn Møsvatn

Lake Kringlesjåen Mjågevatn Tjønndalstjønnan Tuevatni Tuevatni Holmavatnet Honserudvatnet Vollenuttjørnan Midtre Krokavatn Møsvatn Langesjå Mortåtjørni Bukketjørn Reinarvatnet Grunnostjørni Kolsnutgrysline Tuddøltjørn Landsetvatnet Middøltjønne Bågåfjelltjørni Hondletjørni Finnerottjørni NVE ID NVE

18854 18894 18910 18919 18923 18954 19079 19228 66946 3 37 11881 12032 12040 12073 12106 12146 12178 12188 12268 12395 12409 Catchm. no. Catchm.

15 15 15 15 15 15 15 15 15 16 16 16 16 16 16 16 16 16 16 16 16 16

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1 97 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 91, 2, 57, 50, 109 Artsdata- banken.no Artsdata- banken.no 109 2, 57, 50, 9, 50 66, 86 50, 86, 100 78, 117 86, 101 78, 117 66, 74 50, 78 86, 97 Artsdata- banken.no 117 117 117 117 117 117 100 50

TOC mg C L C mg TOC

-1

1,6 2,2 0,5

Ca mg L mg Ca

-1

0.94 0.67 2.41 0.64 0.57 0.90 0.75 0.74 0.79 0.42 0.47 1.66 0.50 1.47 0.56 1.33 0.43 0.44 0.51 pH

6.10 6.00 6.80 6.10 6.49 6.44 5.68 6.39 5.80 6.09 6.12 6.50 6.24 6.65 6.29 6.55 6.19 6.02 5.80 Date

12.09.84 15.09.83 12.09.84 12.09.84 02.10.97 21.09.94 13.09.83 21.09.94 15.09.83 21.09.94 15.08.00 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 24.09.95 15.09.83 No. pos. rec. pos. No.

10 7 1 1 1 1 1 1 2 1 1 No. exam. No. 12 0 0 8 1 1 1 1 1 1 3 1 1 0 0 0 0 0 0 0 0 0 6685973 6667777 6668643 6676361 6680450 6676124 6684434 6693400 6681420 6691453 6674870 6689697 6687654 6669784 6698507 6692302 6693404 6691715 6687392 6687524 6684136 6684637 EPSG:25833

123922 133315 131592 123821 134023 113053 106784 110808 119289 113371 115955 110072 109773 131775 117274 128236 116563 125474 118023 101792 118551 114440 P. phoxinus P.

x S. alpinus S.

x x x

Max. depth m depth Max.

Min. depth m depth Min.

Area km Area

2

20.55 0.79 1.10 31.26 2.03 2.68 3.17 2.05 2.81 2.33 1.63 1.35 3.03 0.36 0.16 0.60 0.55 0.38 0.14 0.23 0.40 0.13

m a.s.l. m Altitude Altitude

1121 1064 1069 1087 1174 1115 1246 1229 1207 1228 1120 1294 1324 1078 1161 1197 1136 1182 1278 1261 1354 1303 Reg. X x Catchment Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Grottetjørn

Lake Mår Nedre Grottetjørn Øvre Gøystvatnet Rosjå Vråsjåen Kallungsjåen Eidsjoren Reksjå Hettefjorden Store Saure Kalven Viuvatnet Prestevatnet Skjortetjørnan Austre Søltjørni Kosadalsvatnet Vestre Søltjørni Melrakktjørnane Tangetjørnan Viktjørn Sletteidvatni NVE ID NVE

36 73 74 75 92 93 94 96 103 104 107 108 109 11750 17925 18132 18133 18187 18456 18562 18642 18661 Catchm. no. Catchm.

17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17 17

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1

98 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 101 50, 101 9 101 100 Artsdata- banken.no Artsdata- banken.no Artsdata- banken.no Artsdata- banken.no 50 Artsdata- banken.no Artsdata- banken.no Artsdata- banken.no 50 86, 117 86, 100 47, 117 100 117

TOC mg C L C mg TOC

-1

1,0 2,0 3,8

Ca mg L mg Ca

-1

0.55 0.65 0.84 0.53 0.66 2.69 1.37 4.82 2.01 0.81 1.37 pH

5.76 5.60 6.05 6.25 6.00 6.70 6.76 7.20 6.85 6.19 6.61 Date

13.09.83 15.09.83 13.09.83 21.09.95 12.09.84 15.09.83 21.09.94 12.10.97 21.09.94 29.10.95 21.09.94 No. pos. rec. pos. No.

1 0 1 1 1 0 No. exam. No. 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 6684111 6683971 6681268 6679611 6676249 6673742 6674654 6672967 6672308 6673633 6672530 6671998 6670974 6671824 6704599 6706737 6709879 6710378 6701741 EPSG:25833

114495 112017 134529 122801 109305 134889 124648 134753 135987 113143 132602 133373 133015 131505 125273 129152 130989 133342 126457

P. phoxinus P. S. alpinus S.

x x x x x x x x

Max. depth m depth Max.

Min. depth m depth Min.

Area km Area

2

0.99 0.42 0.42 0.35 0.38 0.11 0.03 0.03 0.52 0.99 0.04 0.17 1.50 1.02 3.68 1.29 0.71 0.05 0.72

m a.s.l. m Altitude Altitude

1302 1306 1178 1155 1220 1160 1200 1119 1110 1147 1089 1082 1082 1080 1117 1101 1187 1195 1181 Reg. Catchment Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Mår/ Gøyst Uvdalselvi Uvdalselvi Uvdalselvi Uvdalselvi Uvdalselvi

Lake Sletteidvatni Sletteidvatni Ljostjørn Bakketjørn Torstjørni Lontjørnane Reinstjørni Ormtjørnane Store Vålåsjå Saure Vesle Langtjørn Nålvatnet Store Nivstjørn Haraldsjå Skarvsvatnet Tøddølvatnet Store Ormetjørni Tjørngrøtjørnane Bollatjørni NVE ID NVE

18679 18698 18721 18845 19044 19068 19072 19104 19121 19131 19132 19152 19162 36649 385 411 427 17543 17773 Catchm. no. Catchm.

17 17 17 17 17 17 17 17 17 17 17 17 17 17 18A 18A 18A 18A 18A

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1

99 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 117 117 117 117 117 110, 86, 75, 115 86,117 115 117 117 100 86, 117 117 115, 86, 75, 117 86 117 117 117 117 117 23, 21, 3, 114, 117 116, 52, 3, 117, 124

TOC mg C L C mg TOC

-1

2,1

Ca mg L mg Ca

-1

1.28 0.66 0.54 1.88 0.42 1.45 0.58 2.78 0.46 1.64 1.20 0.90 1.27 0.65 0.39 0.62 1.59 1.42 1.57 0.64 pH

6.55 6.27 6.08 6.76 5.98 6.68 6.33 6.95 5.97 6.56 6.54 6.35 6.55 6.29 6.15 6.32 6.64 6.63 6.73 6.27 Date

21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 29.10.95 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 No. pos. rec. pos. No.

3 1 0 1 3 1 4 3 No. exam. No. 0 0 0 0 0 4 1 1 0 0 0 1 0 4 1 0 0 0 0 0 5 4 6700934 6700826 6699604 6698172 6698072 6694102 6695185 6698024 6699369 6699829 6699607 6698934 6697263 6693775 6695521 6693752 6694234 6693465 6690674 6688573 6712627 6720601 EPSG:25833

131680 129576 130951 133127 129358 133668 126931 143460 136935 127220 122882 123493 124531 142350 131830 131663 124708 125832 134645 137346 125791 103447 P. phoxinus P.

x x x S. alpinus S.

x x Max. depth m depth Max.

15,5 Min. depth m depth Min.

4,3

Area km Area

2

0.17 0.86 0.48 0.29 0.20 1.22 2.37 0.44 0.25 0.13 0.11 0.42 0.23 12.53 0.11 0.27 0.28 0.25 0.19 0.39 2.37 1.98

m a.s.l. m Altitude Altitude

1137 1184 1149 1206 1213 1064 1172 1223 1205 1271 1190 1190 1172 1060 1103 1102 1198 1199 1217 1234 951 1148 Reg. x Catchment Uvdalselvi Uvdalselvi Uvdalselvi Uvdalselvi Uvdalselvi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Ølmosåi Lågen Lågen

. Halstjørnane.

Lake Kruketjørna Fiskeløktjørni Langetjørni Gryttjørni Gråvårtjørn Vikvatn Smågefjorden Damtjørn Mjågesjoren Falkenuttjørnane Ø Halstjørn Dragøytjørni Sønstevatnet Kvonneviertjørni Afdalstjørni N. N. Holstadtjørni S. Holstadtjørni Nutetjørnane N. Gryttjørni Store Orsjoren Veslekrækkja NVE ID NVE

17790 17796 17823 17868 17887 409 410 426 17810 17837 17848 17867 17935 17949 17979 18052 18055 18089 18191 18264 390 391 Catchm. no. Catchm.

18A 18A 18A 18A 18A 18B 18B 18B 18B 18B 18B 18B 18B 18B 18B 18B 18B 18B 18B 18B 19A 19A

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1 100 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 116, 117 123 116, 79, 3, 23, 124 84, 6, 23, 100, 108, 109, 113 28, 34, 117 117, 116, 3, 124 72, 52, 2, 3, 116, 94, 109, 117, 124 3, 23, 117 117 65, 7, 5, 4, 108, 113, 117 83, 79, 25, 100 86 83, 79, 25, 82, 86, 79, 126 94,26, 117 43, 41, 14, 46, 44, 45, 118 44, 43, 41, 118 46, 79,

TOC mg C L C mg TOC

-1

1,6 1,0

Ca mg L mg Ca

-1

0.48 0.65 1.32 1.56 1.18 1.11 1.81 1.30 1.29 1.38 3.03 2.40 2.53 2.59 0.28 0.35 pH

6.09 5.49 6.90 6.61 6.52 6.56 6.72 6.51 6.59 6.55 6.93 6.87 6.80 6.85 5.74 5.69 Date

21.09.94 31.05.93 22.07.67 29.10.95 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 02.10.97 30.07.98 11.07.78 21.09.94 25.07.93 25.07.93 No. pos. rec. pos. No.

1 9 3 7 0 2 12 3 7 3 5 3 1 8 4 No. exam. No. 1 13 4 8 2 4 15 4 0 7 3 7 5 3 15 14 6723196 6725710 6706544 6698934 6695177 6714906 6715697 6700686 6699586 6692522 6693471 6697139 6690862 6712322 6730136 6728768 EPSG:25833

101435 98419 113050 97530 105307 106335 101185 116530 111939 93966 85395 79991 75845 96915 99168 99538 P. phoxinus P.

x x x x

S. alpinus S. Max. depth m depth Max.

26,5 42,0 22,8 16,0 11,8 18,0 28,0 Min. depth m depth Min.

8,0 10,0 7,0 4,0 2,8 4,8 6,9

Area km Area

2

4.18 3.33 3.22 11.04 1.55 6.33 13.70 5.09 1.69 18.38 3.12 10.88 1.70 2.95 0.40 1.13

m a.s.l. m Altitude Altitude

1151 1180 1112 1210 1197 1113 1130 1158 1165 1223 1243 1244 1334 1155 1237 1233 Reg. Catchment Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen

Lake Storekrekkja Dragøyfjorden Geitsjøen Langesjøen Geitvatnet Øvre Hein Skrykken Krossvatnet Lakjen Nordmannslågen Dimmedalsvatnet Skaupsjøen Svartevasstjørni Svartevatnet NVE ID NVE

392 393 394 395 396 414 415 416 417 418 419 420 421 428 17018 17040 Catchm. no. Catchm.

19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1 101 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 23 40 117, 40 40 23, 24 40 3, 117 117, 116, 3, 124 79 40 117 109, 3, 100 3, 117 43, 44 100 117 116, 52, 3, 124 124 3 3 117, 86, 3, 124 117 117

TOC mg C L C mg TOC

-1

2,8 2,7

Ca mg L mg Ca

-1

1.47 2.13 0.80 3.12 5.01 1.88 4.47 1.06 1.50 1.76 2.96 1.97 1.00 pH

6.66 6.84 6.34 6.97 7.21 6.80 7.14 6.57 6.61 6.96 6.68 6.39 Date

21.09.94 21.09.94 21.09.94 21.09.94 12.10.97 21.09.94 12.10.97 21.09.94 26.07.92 14.07.67 21.09.94 21.09.94 21.09.94 No. pos. rec. pos. No.

1 2 4 1 0 3 1 7 1 0 0 0 1 No. exam. No. 1 0 0 0 2 0 4 2 1 0 3 0 1 7 0 0 5 1 1 2 3 0 0 6720445 6720331 6721178 6719802 6719115 6718774 6717924 6718724 6718971 6716573 6715599 6715696 6714827 6716874 6714372 6712469 6710330 6711896 6710045 6710191 6709350 6710074 6706344 EPSG:25833

114730 117391 102267 117501 118578 117084 120435 105066 96712 122037 118134 114588 122190 90493 111175 99207 113349 102161 119782 109866 107988 100267 120711 P. phoxinus P.

x x x x x x x x x x x S. alpinus S.

x x x x x x x

Max. depth m depth Max.

Min. depth m depth Min.

Area km Area

2

0.31 0.02 0.71 0.11 1.00 0.26 0.84 0.37 0.22 0.15 0.52 0.18 0.43 0.82 0.10 0.28 1.23 0.17 0.08 0.10 0.49 0.34 1.70

m a.s.l. m Altitude Altitude

1201 1167 1157 1164 1162 1163 1161 1138 1162 1156 1121 1183 1079 1236 1180 1225 1075 1150 1022 1128 1135 1157 1123 Reg. Catchment Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen

Lake Langetjørni Sandtjørni Båtstjørni Urdetjørni Breidvatnet Korta Nedre Bjørkevatn Heinumgen Hetjørni Raudtjørn Bjordalsvatn Herbutjørnan Orsjøtjørni Skiftesjøen Grønenuttjørnan Lometjørni Nedre Hein Skaupungen Halstjørni SelstjørniVesle SelstjørniStore Skaupsjøtjørnan Nordskarvet NVE ID NVE

17318 17320 17332 17336 17343 17364 17366 17397 17411 17422 17454 17456 17460 17477 17492 17565 17583 17584 17591 17612 17617 17624 17653 Catchm. no. Catchm.

19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1

102 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 86, 117 117 86 86 117 117 84, 117 86, 117 86, 83, 79, 100 86, 117 84, 117 79 86 117 117 86, 117 117 100 25 117 108 117 83 79, 25, 100 117

TOC mg C L C mg TOC

-1

1,0 3,5 1,7

Ca mg L mg Ca

-1

2.40 1.71 1.77 2.03 1.10 3.30 1.20 1.74 1.52 1.44 2,17 1.61 2.03 2.90 1.32 1.04 2.85 0.55 0.83 pH

6.84 6.72 6.60 6.76 6.35 6.52 6.51 6.66 6.68 6.57 6,77 6.59 6.74 6.84 6.55 6.35 6.65 6.37 6.30 Date

21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 29.10.95 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 21.09.94 02.10.97 30.07.98 21.09.94 21.09.94 24.07.01 12.09.97 21.09.94 No. pos. rec. pos. No.

1 1 1 2 1 3 1 0 1 1 1 1 1 No. exam. No. 1 0 1 1 0 0 2 1 3 1 2 1 1 0 0 1 0 0 0 0 1 0 2 0 0 6707809 6708108 6705959 6705361 6701694 6703502 6702249 6702002 6701916 6700797 6699604 6700892 6699618 6698601 6698337 6697238 6696625 6695359 6697360 6695944 6695153 6693651 6695803 6692928 6692561 EPSG:25833

100511 99694 94207 97434 112359 94866 102363 93717 78918 92044 105566 80443 92249 93030 95448 90270 98330 96849 85109 94612 89462 98972 75958 107376 106117 P. phoxinus P.

x x x x

S. alpinus S.

Max. depth m depth Max.

Min. depth m depth Min.

Area km Area

2

0.43 0.06 0.06 0.77 0.88 0.60 0.77 1.63 1.02 0.44 0.26 0.10 0.13 0.59 0.21 1.13 0.63 0.69 0.06 0.21 0.74 0.54 0.21 0.27 0.10

m a.s.l. m Altitude Altitude

1170 1170 1212 1207 1167 1213 1209 1211 1256 1214 1209 1260 1222 1220 1210 1235 1220 1220 1294 1237 1225 1223 1346 1230 1240 Reg. Catchment Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen

. Krakavadtjørn

Lake Kalvetjørnan Kalvetjørnan Stigstuvtjørni Holmetjørnane Dagfisketjørni A Bakkatjørn Flotatjørni BakketjørnV. Nordvatnet Kleivshovdtjørn Langesjøtjørni Holmavatn Skrovstjørni Brutjørnan Lomavikan Storfisktjørni Torjustjørnan Torjustjørnan Reinsmyrtjørni Veisetjørnan Ø Nottartjørni Gravagjelstjønn Feitfisktjørnan Bakketjørnan NVE ID NVE

17682 17685 17746 17750 17800 17807 17826 17843 17898 17899 17908 17959 17965 17981 17999 18023 18050 18076 18086 18096 18164 18199 18208 18211 18239 Catchm. no. Catchm.

19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1

103 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 79,100 25 83 25 83 25, 79 117 82, 86 79, 25, 100 117 117 100 100 25 25 124 23, 117 100 117 86 117 23 22 3, 23, 117 3

TOC mg C L C mg TOC

-1

0,4 1,6 0,6 1,3 0,4

Ca mg L mg Ca

-1

2.84 2.04 2.13 0.37 0.52 1.11 0.51 3.95 2.03 2.65 1.90 0.80 1.31 4.37 1.46 3.52 2.47 3.31 pH

7.07 6.56 6.32 5.90 6.01 6.00 6.20 6.44 6.21 6.82 6.44 6.83 6.93 6.60 6.55 7.00 6.63 6.62 6.90 7.11 Date

02.10.97 17.07.96 17.07.96 28.07.95 21.09.94 28.07.95 21.09.95 21.09.94 21.09.94 02.10.97 02.10.97 31.07.03 31.07.03 17.07.67 21.09.94 09.10.97 21.09.94 21.09.94 21.09.94 21.09.94 No. pos. rec. pos. No.

0 1 0 1 0 4 1 1 1 1 3 3 1 No. exam. No. 1 0 1 1 1 1 0 7 0 0 0 0 0 0 0 2 1 0 0 1 0 1 3 3 1 6694635 6694619 6694381 6693954 6693986 6692957 6690638 6692150 6689092 6688826 6688587 6690318 6688873 6688201 6687575 6712787 6703571 6705286 6701192 6702161 6722322 6722372 6722207 6720539 6719724 EPSG:25833

77497 76469 75897 74515 75917 76839 105369 74729 102718 87396 98746 82119 88840 81147 82385 109162 113891 76272 110914 93632 121472 122888 117562 119750 124886 P. phoxinus P.

x S. alpinus S.

x Max. depth m depth Max.

28,0 Min. depth m depth Min.

8,9

Area km Area

2

0.30 0.06 0.04 0.23 0.04 0.16 0.87 1.88 0.19 0.47 0.58 0.15 0.14 0.14 0.21 0.58 0.47 0.12 0.25 1.29 0.19 0.04 0.20 1.43 0.21

m a.s.l. m Altitude Altitude

1303 1313 1326 1357 1330 1325 1307 1331 1255 1246 1225 1268 1235 1285 1314 1112 1157 1308 1166 1212 1265 1237 1236 1183 1129 Reg. Catchment Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Lågen Ufysja Ufysja Ufysja Ufysja Ufysja Bessvatnet

Lake Bessvatnet Øvre X-tjørn Høgevardetjørn Y-tjør n Dimmedalstjørni Steintjørni Bismarvatnet Vegarhovdtjørna Langebutjørnan Sørtjørn Lakadalstjørnane Langebutjørnan Lakadalstjørnane Holbergtjørnane Heintjønne Hølen Andersosen Danemannstjønni BakketjørnV. Tindevatnet Tobakkstjørni Tuvetjørni Holværvatnet Svantjørni NVE ID NVE

18257 18274 18289 18290 18305 18336 18338 18374 18445 18466 18469 18478 18536 18612 18650 66935 66954 67633 276916 279619 17222 17224 17244 17281 17297 Catchm. no. Catchm.

19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19A 19B 19B 19B 19B 19B

. Continued. Region C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1

104 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019

References 109, 23, 3, 117 30, 29, 13, 89 33, 31, 31 30, 29, Own data Own data Own data Own data Own data Own data 37, 36, 35, 81 39, 38, Own data Own data Own data 29,30,89

TOC mg C L C mg TOC

-1

Ca mg L mg Ca

-1

1.83 1.80 0.70 2.30 pH

6.79 6.53 6.43 6.77 Date

21.09.94 26.08.05 28.08.05 23.08.05 No. pos. rec. pos. No.

6 6 2 8 3 No. exam. No. 6 9 3 0 0 0 0 0 0 9 0 0 0 3 6716355 6725146 6727693 6721906 6720705 6720862 6720646 6720017 6719472 6718885 6719422 6718655 6718009 6723166 Institute Freshwater of Research, Drottningholm. 183-201. 1969: Report 49, No EPSG:25833

125104 103584 103408 107583 108533 111069 112980 109068 112034 113282 109729 115187 111304 106124 NIVA Rapport L.NR. 2387-1990. 45 pp. (In pp. Rapport 45 Norwegian). NIVA L.NR. 2387-1990. P. phoxinus P.

x x x x x x x x x x x x x

S. alpinus S. Max. depth m depth Max.

6,0 Min. depth m depth Min.

1,7

Area km Area

2

0.90 9.44 0.48 0.15 0.19 0.12 0.02 0.07 0.13 1.57 0.35 0.10 1.21 1.74

m a.s.l. m Altitude Altitude

1182 1147 1149 1162 1169 1220 1240 1176 1193 1192 1180 1197 1190 1147 Reg. x x x Catchment Ufysja Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Ørteråni Trestiklan

Lake Ljosevatnet Ørteren Øvre Veslevatnet Lonin Håkonstjørnane Grønehølen Steintjørni Urdavatnet Skjerjavatnet Grønehølen Underhaugtjørn Grønevatnet Lægreidvatnet NVE ID NVE

17401 553 17104 17283 17312 17319 17327 17356 17363 17371 17372 17381 17395 67975 Catchm. no. Catchm.

19B 20 20 20 20 20 20 20 20 20 20 20 20 20

. Continued. Region fiskeribiologiske undersøkelser LFI Uni Research i perioden LFI Miljø, Rapport 60 (In pp. 1999-2013. Norwegian). 235. nr. (In pp. 45 Norwegian). Universitetet i Bergen. Rapport 111-2000. nr. (In pp. 120-2002. 39 Norwegian). C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E C/E Appendix 1 REFERENCES Aanes KJ,1. Resipientundersøkelser Brettum Holtan 1990. G. P, for kommune i Isdalsvann med Isdøla. Crustacea, 1969. especially2. Aass P. Lepidurus arcticus Pallas, as brown trout food in Norwegian mountain reservoirs. Fiskeribiologiske undersøkelser Fiskerikonsulenten Amundsen (In Dagalivassdraget3. 1976. pp. for 285 Rapport Øst-Norge. Norwegian). for T. IV-3.2. 1974-75. Na Gabrielsen Barlaup 4. S-E, BT, Halvorsen Skoglund GA, A, Hobæk H, Lehman Wiers T, Pulg SkårGB, U, B,Normann Bjornesfjorden E, Restaurering– Fjellheim A. 2014. gyteområder av og Kleiven E, Raddum Barlaup BT, 5. GG, Gabrielsen S-E, Johannessen undersøkelser A. 2000. Fiskebiologiske i Bjornesfjorden, Laboratoriet august ferskvannsøkologi 1999. for innlandsfiske og (LFI), Kleiven E, Skoglund6 Barlaup undersøkelser H. 2002. BT, Fiskebiologiske i Langesjøen, Laboratoriet august 2001. ferskvannsøkologi for innlandsfiske og (LFI), Universitetet i Bergen. Rapport nr.

105 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019 Norwegian 101-112. Journal Zoologi 21: of sumartemperatur interaksjonar og innan bestandane er avgjerande faktorar. Norges landbrukshøgskole, Institutt ) biologi og utbredelse og ) biologi Særtrykk i Norge. (In Fiskerforenings pp. og 32 Norwegian). Norges Jæger- av Tidsskrift 1915. , Lægreidvatn,, Øvre Nedreog Trestiklan, kommune Fylkesmannen Hol 1994. Buskerud,i miljøvernavdelingen. Rapport nr. L) i en vest-øst gradient L) på Hardangervidda. Universitetet miljø biovitenskap, for og Institutt naturforvaltning. for Master thesis. på Vestvidda: Snø, Ørteren Notat. 7 pp (In Norwegian). Gammarus pulex , Lægreidvatn, Øvre Nedre og Trestiklan, miljøundersøkelser. og kommune Hol Fisk- 2005. Rapport. (In pp. 36 Norwegian). Pallas, i regulerte vann. egg av i reguleringssonen Forekomst klekking og egg. Laboratoriet av ferskvannsøkologi for innlandsfiske (LFI), Salmo trutta Ørteren Lepidurus arcticus Fiskerapport nr. 8-1998. 17 pp. (In pp. Norwegian). Fiskerapport 17 8-1998. nr. Prøvefiskerapport frå Tinnhølen, 02.08.2000. Eidfjord fjellstyre. (In 2 pp. Norwegian). . Continued. Bjornesfjorden Laboratoriet ferskvannsøkologi 1999-2007. for innlandsfiske og (LFI), Universitetet i Bergen. Rapport (In pp. 70 Norwegian). 150-2008. nr. (In pp. 43 Norwegian). (In pp. 33 Norwegian). Universitetet Rapport i Oslo. 140-1993. nr. naturforvaltning. og biologi for Fiskerapport (In 20 pp. Norwegian). 13. nr. naturforvaltning. (In pp. 12 Norwegian). Universitetet Rapport i Oslo. 22-1975. nr. ferskvannsøkologi innlandsfiske og (LFI), Universitetet Rapport (In pp. i Oslo. 36 Norwegian). 90-1986. nr. (In pp. Norwegian). 10 3-2010. nr. (Inpp. Norwegian). Norwegian). (In (In pp. 33 Norwegian). 14-1995. Appendix 1 Sandven OR, Skoglund Barlaup H, Gabrielsen BT, 7. S-E, Kleiven E, Lehmann Wiers T, G, Fjellheim Å. A, 2008. Halvorsen Restaurering GA, A, Hobæk Tysse gyteområder av prøvefiske og i Bjørkhaug8. E, Hamre hos ørretunger Vekst 2005. B. ( Bjørtuft9. Fiskeribiologiske undersøkelser SK, 1993. Saltveit i forbindelse SJ. med planlagte overføringer til Mår kraftverk Laboratoriet i Telemark. ferskvannsøkologi for innlandsfiske og (LFI), Borgstrøm10. R, Dokk 2000. JG. Rekruttering til aurebestandar Borgstrøm R, Sporan11. NR. Prøvefiske 1998. i Ambjørnsvatn, Statsallmenning, Bestandsanalyse i 1997. forslag til og driftsplan. Norges landbrukshøgskole, Institutt og biologi for Borgstrøm The12. effect increased of R. 1973. fluctuation level upon the brown trout population Mårvatn, of a Norwegian reservoir. Skjoldkreps, Borgstrøm13. R. 1975. Borgstrøm Aurebetanden R.14. 1990. i Svartavasstjønn. Notat. (In Norwegian). Borgstrøm Midtre R.15. 2003. Grøndalsvatn Aurebestandar 2003. i Ullensvang statsallmenning, (In Faktaark2 pp. Norwegian). 1-2003. nr. 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Jæger- Sømme Tidsskrift:21. 1933. ID. 297-301. En studie grundaatens over 22. Dahl K. 1915. eller matfloens ( Studier forsøk Ørret og over Ørretvand. og Dahl23. K. 1917. Kristiania. Det Mallingske Bogtrykkeri. pp (In Norwegian). 182 Dahl24. Studier K. røje i ørretvand. 1920. over Fiskerforenings og Norges Jæger- Tidsskrift (In Norwegian). 233-248. 1920: Eidfjord Diverse fiskerapporter.25. fjellstyre www.eidfjordfjellstyre.no 1995-2010. (In Norwegian). Å. 2004. Elnan26. Fiskeundersøkjingar G, Tysse i Skaupsjøen. Elnan 2000. G. 27. Enerud28. Fiskeribiologiske undersøkelser Garnås J, i Geitvatnet, E. 1987. Uvdal kommune, Nore og Buskerud fylke. Fylkesmannen i Buskerud,pp. 17 miljøvernavdelingen. 1987. Rapport F-1, nr. Enerud Garnås 29. J, Fiskeribiologiskeundersøkelser E. 1995. i Enerud30. 2006. J. Fiskeribiologiske undersøkelser i Eriksen Garnås SD, 31. Prøvefiske E. 1986a. driftsplan og Øvre for Trestiklan Prosjekt Fjellfisk, 1986. (In 9 pp. rapport Norwegian). nr.19/86. 106 Qvenild and Hesthagen: Lepidurus arcticus on Hardangervidda, Norway Fauna norvegica 39: 77–110. 2019 Hydrobiologia 73-84. 519: . . Innlandsfiske. LFI-UNIFOB. Universitetet i Bergen. Rapport Økol på Hardangervidda. Norges landbrukshøgskole, Institutt og biologi for Lepidurus arcticus L., i innsjøer L., på Hardangervidda. Norges landbrukshøgskole, Institutt biologi for L., i Krokavatn L., Salmo trutta Salmo trutta Rapport fra Høyfjellsøkologisk pp. 17 Forskningssenter, (3). Finse, 1973 Norge. L. i ulikt tette bestandar, radiotelemetristudium eit i tre vatn på Hardangervidda. Norges landbrukshøgskole, Institutt for i forhold til i forhold tettheten ørret, av 1987. Prosjekt (In 8 pp. Fjellfisk, Norwegian). rapport 1987. nr.91/87. Salmo trutta Ørteren Daphnia longispina Circumpolar analysis morphological of and genetic diversity in the Notostracan thesis. 46 (In pp. Norwegian). Rapport (In pp. Norwegian). 35 2/ nr. 86. . 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