Faunal Finds from Alpine Ice: Natural Or Archaeological Depositions?

JGA 3.1 (2016) 79–108 Journal of Glacial Archaeology ISSN (print) 2050-3393 https://doi.org/10.1558/jga.32414 Journal of Glacial Archaeology ISSN (print) 2050-3407

Faunal Finds from Alpine Ice: Natural or Archaeological Depositions?

Jørgen Rosvold

University Museum, Norwegian University of Science and Technology

[email protected]

Inland ice covers large areas of the world’s surface, but the ecology of the ice itself is poorly studied and largely unknown. A large varietyof melted out faunal finds from glaciers and ice patches around the world have been discovered for more than 150 years. These finds hold a unique and largely untapped information potential for archaeology, faunal history and glacial ecology. In order to retrieve information from this frozen databank we need a better understanding of how the material were deposited. This article provides a background for glacial faunal finds worldwide and presents the relatively large Norwegian collections for the first time. The Norwegian finds are very well preserved, allowing good insights into the taphonomy of the finds. While most finds seem to be naturally deposited, many of the sites are interesting hybrids between archaeological and natural history sites. Potential implications and prospects for future management and research are further discussed.

Introduction Despite covering about 10% of the Earth’s land areas (Zemp et al. 2008), inland ice remains one of the least understood ecosystems in the world and ice covered landscapes are usually considered as barren wastelands, devoid of life. The discoveries of diverse microbiological communities of ice-dwelling viruses, bacteria, algae and fungi on and in the ice, are currently changing this view of the ice among biologists (Hodson et al. 2008; Anesio and Laybourn-Parry 2012). Some of these micro-organisms have even been shown to be primary producers that provide a nutrient foundation for distinct glacial ecosystems (e.g. Anesio et al. 2009), but many of the nutrients in these systems are deposited through windblown (aeolian) material in the form of arthropods, mineral dust, pollen and other small plant materials (Kaisila 1952; Wilson 1958;

Keywords: Alpine, birds, butchery, glacier, glacial melting, gnaw marks, Holocene, ice, mammals, taphonomy

© Equinox Publishing Ltd. 2018, Office 415, The Workstation, 15 Paternoster Row, Sheffield, S1 2BX 80 Jørgen Rosvold Edwards and Banko 1976; Swan 1992). This windblown material offers easily available food for e.g. alpine spiders and birds that use the ice as periodic foraging habitats (Edwards 1987; Rosvold 2016). The glacial ecosystems even include many larger birds and mammals, and several vertebrate species are known to actively use ice covered land for various activities on a seasonal basis (Rosvold 2016). However, the diversity and function of vertebrates in these ecosystems are poorly understood and studied. As the inland ice continues to melt away due to a warmer climate we risk losing most of these poorly understood ecosystems within a short time (Vaughan et al. 2013). In regards to this, glacial research is currently contributing invaluable data for understanding and appreciating the ecological history and importance of alpine ice. A positive side-effect of global warming is that the melting ice is revealing unique traces of its own cultural and ecological history, through finds of melted out organic materials that have been preserved in the ice for up to thousands of years. Archaeolo- gists have been aware of these excellent repositories of long-term historical material for some time and have, through a variety of finds that have emerged from the ice, documented that humans have been important elements of these icy ecosystems for many millennia (Dixon, Manley and Lee 2005; Reckin 2013; Dixon et al. 2014). Human use of the ice is varied, but the recovered archaeological material indicates that the alpine ice has been used as hunting grounds in many areas around the world, in particular for the hunting of alpine ungulates (Dixon et al. 2014). Numerous faunal finds have been made in connections with glacial archaeological investigations, but they generally tend to receive much less attention than the spectacular archaeological finds. Thus, they have escaped the focus of most biologists and the literature is usually scarce concerning such finds. However, these faunal remains have a great

Figure 1 Worldwide distribution of published faunal finds from glaciers and alpine ice patches. White areas represent ice covered land and stars represent mountain areas where finds have been made. Each star may thus represent several sites within a larger area. See Appendix 1 for more details and references. Background map is modified from Natural Earth maps.

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 81 potential to enhance the ecological significance and value of ice-covered landscapes, and to provide key information for interpreting the context of these glacial sites. There is, however, a need for a better understanding of what the faunal material actually reflects and especially how the finds were deposited in the ice. Given the history of human interactions with many glaciated areas, it is possible that large numbers of the faunal material could be a result of hunting activities, but to verify this assumption it is necessary to link faunal remains from these sites to human activities. From an ecological perspective, it has also been assumed that these glacial landscapes represent unique archives of alpine natural history (Rosvold 2016), which attract animals from a wider area and acts as a form of natural “pit trap,” with a more biologically representative faunal collection than is regularly found on archaeological sites. This depends on how representative the material actually is. Here I will discuss these issues and reflect upon the deposition of vertebrate faunal material in the ice. The focus for the discussion is on what the diversity of finds, dates and taphonomic conditions can tell us about the nature of the material and its research potential. I only include faunal remains that have been found in connection with superficial land ice, i.e. glaciers and ice patches. Ice patches (often called snow patches) are defined as smaller perennial accumulations of ice and snow, which in contrast to glaciers have little or no ice flow (Fujita et al. 2010; Serrano et al. 2011). Invertebrate animals (e.g. Lockwood et al. 1991), and obvious human artefacts made out of animal parts, like bone arrowheads (e.g. Callanan 2014), arrow fletchings (e.g. Dove, Hare, and Heacker 2005) or clothing (e.g. O’Sullivan et al. 2016) are not included here. Nor are finds from permafrost (e.g. Boeskorovet al. 2014) or cave ice (e.g. Lauriol et al. 2001) included, which are separate ecological phenomena. I start by providing a background of reported finds from glaciers and ice patches worldwide, before giving a first presentation of the relatively large and hitherto unreported Norwegian faunal finds. The Norwegian glacial faunal materials are generally very well preserved and allow good insights into the taphonomy of the finds.

Background An overview of reported vertebrate faunal remains found in connection with superficial land ice worldwide are presented in Figure 1 and Appendix 1. Reports of such finds are often obscure and presented as short notes or as unpublished reports in local languages. It is likely that there are additional finds reported in grey literature or not reported at all. Nevertheless, observations of animal remains on or in alpine ice have been recorded at least since the late nineteenth century “golden age of moun- taineering” and other alpine expeditions (e.g. Tyndall 1871, 331; Meyer 1891). One of the more renowned finds is that of a mummified leopard (Panthera pardus) carcass found on a glacier on Mt. Kilimanjaro (Latham 1926), which is reported to have in- spired Hemmingway’s “The Snows of Kilimanjaro”. These finds were just considered curiosities, but possibly the first reflection on the potential old age of such melted out animal remains was made by the geologist Peter A. Øyen in 1913. He reported a local farmer’s observation of some unusually large animal bones at the edge of a melted ice patch near Digervarden in south-central Norway. The bones were never

© Equinox Publishing Ltd. 2018 82 Jørgen Rosvold recovered due to a following snowfall and colder climate, but were speculated to be of a mammoth (Øyen 1913). Subsequent discoveries of archaeological artefacts melting out of the ice in Norway in 1914 stimulated the occasional collection of nearby animal bones throughout the twentieth century (e.g. Farbregd 1972; Callanan 2012). However, systematic collections and searches for glacial faunal remains did not be- gin before the discovery of large amounts of melted out reindeer (Rangifer tarandus) remains along with a 4000-year-old hunting dart in the Yukon mountains in 1997 (Kuzyk et al. 1999). This discovery stimulated several large-scale multidisciplinary investigations of alpine ice patches in North America (Farnell et al. 2004; Hare et al. 2004; Dixon, Manley, and Lee 2005) and Norway (Callanan 2012; Nesje et al. 2012), which are starting to provide rich and varied collections of faunal remains. Melted out faunal remains from glaciers and alpine ice patches have been found in large parts of the alpine world, from the arctic and even to tropical mountains (Figure 1; Appendix 1). Including the Scandinavian finds reported here (see below), melted out remains of at least 68 species of mammals, birds and even fish have been reported (Appendix 1). This is far more than the number of species that has been scientifically described to actively use inland ice for various activities (Rosvold 2016). Thus, these finds can prove valuable for improving our understanding of glacial ecology and the history of use of the inland ice by various organisms. Large-scale systematic surveys for such faunal material have only been done in North America and Scandinavia, and there are some very apparent data gaps compared to the distribution of glaciated areas. For some mountainous areas like Antarctica, New Zealand and the Andes, the lack of finds can partly be explained through the absence of large, herd-living, alpine ungulates that often visit such areas en masse, like reindeer. However, other mountainous areas that might be expected to have potential for a variety of faunal finds, like Siberia, the Himalayas and the Andes, are also generally lacking in finds, likely due to lack of sampling and/or publication. If effort is put on searching for and documenting melted out faunal finds, it is likely that many more finds will be made worldwide. Numerically, the great majority of finds represent alpine ungulates; in particular reindeer, Dall sheep (Ovis dalli), bighorn sheep (Ovis canadensis) and American bison (Bison bison) (Farnell et al. 2004; Hare et al. 2004; Dixon, Manley, and Lee 2005; Andrews, MacKay, and Andrew 2012; Lee 2012). Collectively, in large find assemblages, these species typically constitute between 70–80% of the faunal finds. These, and most of the other reported finds, are of alpine (or seasonally alpine) species, of which some are scientifically documented to visit glaciers and ice patches for various activities, particularly during summer (Rosvold 2016). A surprisingly large variety of finds, however, is of species that are not associated with alpine ecosystems. A number of mummified tropical lowland birds found on small glaciers at the Carstensz Mountains in Indonesia are likely animals that died while trying to fly across the mountains or were carried up by strong updrafts (Schodde et al. 1975). Some of the African finds, like the mummified leopards (Pan- thera pardus), African civet (Civettictis civetta) and African wild dog (Lyacon pictus) are more cryptic and might just be stray animals (Latham 1926; Guest and Leedal 1954;

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 83 Mizuno 2005). Other finds have evidently been brought up to the sites by other organisms. This is particularly obvious with fish remains that were found on two sites in North America (Beattie et al. 2000; Dixon, Manley and Lee 2005). Fish do not live on glaciers or ice patches and thus these finds are probably human food remains, which is also highly likely for the remains of a chum salmon (Oncorhynchus keta) found close to the human body of the Kwäday Dän Ts’ìnchi discovery (Beattie et al. 2000), or they could be stomach contents from birds or mammals that have died on the sites. Finds of mule bones, at the Theodul Glacier in the Alps, have likewise been interpreted to belong to a domestic animal that died while crossing a mountain pass with humans (Savioz 2015). Generally, the depositional events of most of the finds are not well known. Are they remains of animals that died on the ice due to natural causes or are they remains from hunting/storage or butchery waste? Apart from the North American and Scandinavian (see below) finds, most of the glacial faunal remains that have been reported are of more or less complete animals or antlers and horns. And aside from the mule bones (see above), none of these reported finds have been directly linked to human activities on the ice. Among the more systematically collected North Ameri- can faunal finds, researchers have noted the general lack of cut marks and evidence for butchery or human modification of bones, but with frequent signs of carnivore gnaw marks (VanderHoek, Tedor and McMahan 2007; Andrews, MacKay and Andrew 2012; Hare et al. 2012). These interpretations have been hampered due to a typically highly weathered surface of the bones. Cut marks are often hard to identify accu- rately, even on well-preserved surfaces, and a coarse surface texture makes spotting potential signs of human interaction even more difficult. Below, I will explore how the Norwegian finds fit the patterns from other known finds and how they might shed more light on the issues above.

Norwegian faunal finds from glaciers and ice patches Faunal finds from glaciers and ice patches in Norway have been collected sporadically since 1914 (Farbregd 1972), largely by local hunters and hikers. Systematic collections of animal remains did not start before 2013. Since then all types of faunal remains have been collected by myself or through an extensive network of researchers, management institutions and local hikers. Here I give a first and broad presentation of the Norwegian faunal finds that I have personally studied (status Dec. 2016), including all the material from my own projects in central Norway (Møre and Romsdal, Nord- and Sør-Trøndelag, and Nordland counties) and most of the faunal material collected by the Oppland County Council in southern Norway. I also include one Swedish find made by Västerbotten Museum, in Västerbotten County, close to the Norwegian border. This includes 1013 individual specimens from 52 different sites (Figure 2). Of these animal remains, 144 samples have been radiocarbon dated. The dates were calibrated into calendar years before present (BP) using CALIB 7.0.2, and the dataset IntCal13 (Reimer et al. 2016). Unless otherwise stated, all dates in the text are in calibrated years BP. A detailed taphonomic analysis of the material is beyond the scope of this paper, but the finds were examined for some general taphonomic patterns in order to look

© Equinox Publishing Ltd. 2018 84 Jørgen Rosvold for indications of depositional events. The material was inspected for cut-marks or other signs of butchery or human modifications, as well as for carnivore gnaw- marks. Further, the general state of preservation was noted, with regards to the presence of soft tissue, adipocere and mineral staining (i.e. vivianite), as well as bone fragmentation and weathering. The central Norwegian material was studied in more detail in regards to the surface preservation of bones, which was visually inspected and classified into six weathering stages (WS), 0–5 where 5 is the most weathered

Figure 2 Map of Norway showing the distribution of glaciers and alpine ice patches with faunal finds marked by stars. The encircled sites are the central Scandinavian sites studied in more detail for surface preservation.

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 85 (Behrensmeyer 1978). The WS of the samples were correlated with rounded calibrated radiocarbon dates using simple linear regression. The results are discussed in comparison with personal observations made on the Yukon faunal material (see Hare et al. 2012) in 2016.

Diversity and age of the faunal remains The Norwegian glacial faunal finds are varied in species composition (Figure 3), body part representation and age (Figure 4, Appendix 2). Faunal remains of at least 20 species have been found thus far. In line with the large North American finds, a large alpine ungulate, i.e. reindeer, dominate the material and constitute 76% of the faunal finds, followed by ptarmigan (all are likely Lagopus muta) and small rodents. The age of the finds are varied and span about 4700 years in time. While approximately 14% of the finds are recent, the 14C dates show that most finds have been in the ice for considerable time. For reindeer, bones from all parts of the body have been found, but antlers dominate the material. About 43% of the entire Norwegian assemblage consist of reindeer antlers, antler fragments and skull fragments with unshed antlers. About 24% of the entire material have been identified as shed antlers of both males and females, as well as calves. As males shed their antlers in early winter, while calves and females shed theirs during the spring, The shed antlers indicate that reindeer are actually

Figure 3 Percentage distribution of Norwegian animal remains from glaciers and ice patches after refitting fragmented bones, n=766 including 93 (12.1%) unidentified remains.

Figure 4 Number of radiocarbon dates of Norwegian faunal samples in 300-year intervals. present on the sites for large parts of the year and not just during the warmest parts of summer and early fall. As with other glaciated sites worldwide, some of the Norwegian finds are of species that are not normally associated with high alpine areas. Domestic animals, i.e. sheep (Ovis aries), horse (Equus ferrus) and cattle (Bos taurus), have been found at several sites and could potentially be associated with human activities on the ice. How- ever, domestic sheep on outfield summer grazing are frequently encountered near ice patches during fieldwork and are likely to be using the ice to cool down or escape insects. Several species of wild animals normally associated with more forested areas, like moose (Alces alces), western capercaille (Tetrao urogallus) and woodpecker (Dendrocopus sp.) could be stray animals or might also have been brought up to the sites. Interestingly, two of the oldest dated ice patch finds in Norway are of capercaille (Appendix 2) which could potentially be a reflection of higher tree-lines in the past. Even some marine species have been found, i.e. several articulated vertebrae of Atlantic mackerel (Scomber scombrus) and some articulated wing bones of razorbill (Alca torda). The mackerel has evidently been brought up to the ice by another organism, while the razorbill could have died while trying to cross the mountains between the Atlantic Ocean and the Baltic Sea.

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 87 It should be noted that the collected material might not exactly reflect the diversity of what was originally deposited on the ice. Most of the finds acquired or reported through local hikers and hunters are reindeer antlers. The relatively small remains, e.g. of small mammals and birds, have only been found through searches that were specifically targeted at finding animal remains. Large numbers of rodent carcasses are often observed on the surface of some ice patches, but few have been collected thus far due to logistics and the view that these are likely of recent age. Radiocarbon dates on a sub-sample of these naturally mummified rodents show that they vary greatly in age (Appendix 2) and, consequently, such remains can provide highly valuable resources for natural history if they were systematically collected. Larger species and more spectacular finds are thus likely somewhat overrepresent- ed in the collected material. If emphasis and awareness are put on documenting all types of animal remains, the diversity of finds would likely increase and become less skewed towards larger species in the future. This would probably hold for most sites worldwide.

Preservation and general taphonomic patterns The faunal finds from the ice are generally very well preserved. The finds range from more or less complete and fully preserved organisms, to fragmented bones, feathers and pieces of skin and hair. There is generally a noticeable difference in the preservation of small and large organisms. Small birds and mammals are typically represented by mummified remains (e.g. Figure 5a); often with most of the inner organs deteriorated, but sometimes intact. On the other hand, remains of larger animals (e.g. ptarmigans and up) are usually found as clean osteological material with very few traces of external soft tissue. This suggests that there is some microbial activity going on within the ice. Many of the bones, however, still contain marrow and some adipocere (e.g. Figure 5b). This difference in preservation between small and large animals could be an effect of more rapid freezing and burial of smaller bodies, while larger animals have been exposed to weathering and scavengers for a longer time. A lack of skeletonized remains of small vertebrates could also be because these remains are harder to spot in the field. Encrustations of the iron phosphate mineral vivianite are frequently found on bones (e.g. Figure 5c) from ice patches, especially from those sites with large deposits of animal (i.e. reindeer) dung. This often vivid blue staining most commonly occurs on bones that have been found in the melted out and accumulated organic debris along the downslope edge of the ice patches. Vivianite usually occur on waterlogged organic remains in iron rich sediments, often when located in contact with iron artefacts (McGowan and Pragnell 2006; Rothe, Kleeberg, and Hupfer 2016). Similar specimens have also been found in some of the glacial faunal remains from North America (Lee, Benedict, and Lee 2006; Pers. obs. on the Yukon material). While e.g. the deposits of reindeer dung in the ice provide a rich source of phosphate, the source of iron in these settings is unclear. It has been suggested that the needed iron could be derived from the blood of the organism itself (Fisher et al. 2012). The presence of vivianite is a good indication of active metal reducing bacteria (Rothe, Kleeberg, and

© Equinox Publishing Ltd. 2018 88 Jørgen Rosvold Hupfer 2016), which could have implications for the preservation of archaeological metal artefacts in the same contexts. In general, the surface of the bones are very well preserved (e.g. Figure 5d), but antlers and long bones are often fragmented in zones where they bend naturally (e.g. Figure 5e). This fragmentation also occur in finds from ice patches, which have generally stable and non-flowing ice, and is thus likely caused by the pressure ex-

Figure 5 (a) Mummified carcass of a Norway lemming. (b) A reindeer mandible with adipocere (white staining) that has been cracked open by carnivore teeth. (c) Broken off reindeer antler point with nodules of vivianite encrustations indicated by arrow (bottom end has been cut off for analyses). (d) A c. 3100 year old reindeer scapula with carnivore gnaw marks. Tooth punctures indicated by arrows. (e) Re- fitted fragments of a reindeer skull with antlers, found spread out over a c. 500 m2 area by the lower edge of an ice patch. Notice the differences in preservation between the two halves of the cranium. Photo: T. S. Kristiansen/NTNU University Museum. (f) Weathered surface, but very well preserved interior of a bone from a Yukon ice patch.

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 89 erted by the weight of the ice. Due to the excellent preservation, these fragments are usually easy to refit. After refitting, the total specimen count was thus brought down from 1013 to 766 specimens (Figure 3). Several distinctive small fragments of antler tines (Figure 5c) have been found in Norwegian ice patches, similar to tine fragments found on some North American ice patch sites and may represent archaeological antler artefacts known from other contexts (Hare et al. 2004; Lee, Benedict, and Lee 2006; VanderHoek et al. 2012). There is, however, no distinctive use-wear on these antler points and they are most likely a product of natural fragmentation. The central Norwegian glacial faunal material, which have been studied in more detail, are in general extraordinarily well preserved and not much weathered. About 50% of the finds are within WS 0 or 1, and when comparing all the radiocarbon dated finds (and considering mummified remains and skin as WS 0) there is no relationship between WS and the age of a sample (n=130, r2=0.007, p=0,335). Interestingly though, when only considering non-modern osteological reindeer finds, which are more comparable, there is a significant correlation between age and WS (n=83, r2=0.048, p=0,046), with a slight tendency for older finds being better preserved (Figure 6). This pattern might suggest that younger finds melted out of the ice before older finds and have thus been exposed to weathering for a longer time before being collected. There is, however, a wide variation in the surface preservation, as expected from such a

Figure 6 Weathering stages (WS) of radiocarbon dated reindeer bones.

© Equinox Publishing Ltd. 2018 90 Jørgen Rosvold diverse sample of finds and sites. This variation is also apparent within sites (e.g. Figure 5e), which shows that it is impossible to tell the relative age of a sample from its visual appearance alone. When comparing the Norwegian material to the large glacial faunal material from Yukon (e.g. Hare et al. 2012), there are some striking differences in preservation. Compared to the Norwegian material, the Yukon material has more preserved soft tissue, in the form of ligaments and tendons, on the large mammal bones. Also, while the interior of the bones are extraordinarily well preserved, the surface of the Yu- koon bones is typically much more weathered (e.g. Figure 5f). A likely reason for this difference is the difference in climatic conditions. The ice patch study area in Yukon is considered semi-arid (Hare et al. 2012), and the Yukon bones exhibit typical signs of repeated freeze-thaw cycles (Pokines et al. 2016) under relatively dry conditions (Pilloud et al. 2016). Central Norway, on the other hand, is considered relatively moist with most areas receiving more than 500 mm of annual precipitation (Moen 1999). Whereas the cold environment of the ice, with little liquid water, offers perfect conditions for preserving animal remains while in the ice, they will encounter harsh conditions once they melt out, if exposed to repeated fluctuations of wet/ dry conditions with flowing oxygen-rich water (Turner-Walker 2007). Thus such remains will likely disappear relatively quickly once melted from the ice in the moister Norwegian mountains. During such conditions, we will only recover the more recently uncov- ered remains, while bones melted out in drier alpine areas, like Yukon, may survive for a longer time while sustaining surface weathering. The generally low WS values of the Norwegian faunal material is ideal for spotting signs of human or animal interactions with the material. Even so, none of the Nor- wegian faunal remains display ambiguous evidence for any kind of human modifications, apart from 13 reindeer antlers that have been chopped or sawn off (e.g. Figure 7). Three of these have so far been dated “TRa-10208,” “TRa-10212” and “TRa-10216” (Ap- pendix 2), of which two are Viking Age and one is modern. The modern one (Figure 7a) is a male skull where one of the antlers have been removed at the base, while the other 12 finds are loose antler parts. These antlers have either been detached at or near the burr (n=4) (e.g. Figure 7b) or they are discarded parts of the brow, bez or top palms (n=8) (e.g. Figure 7c). The four antlers that have been detached near the burr are small and likely female antlers. A possible explanation might be that they were detached to ease the transportation of carcasses. The discarded palms might be a result of pre-processing of antlers intended for tool production, where only the beams were brought off site. Collection of reindeer antlers as raw material, especially for combs, was an important trade in Norway during the Viking and Middle Ages (Christensen 1986; Mikkelsen 1994; Indrelid and Hufthammer 2011). Antler finds from production sites suggest that entire antlers were collected, but some of the pieces found at the ice patches are consistent with known waste material from such productions (MacGregor 1985; Flodin 1989; Weber 2007). These objects, however, could be related to local use of a different character and more detailed studies and radiocarbon dates are needed to resolve this. It cannot be excluded that chop or saw marks on other bones have been lost through degradation of bone edges. The edges, however, are usually well preserved and no cut-

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 91 marks have been found on the intact parts of the bones where they normally appear on other archaeological material. Several more or less complete bones from all parts of the body have been found, but none shows any signs of butchery. While evidence of human interaction with the melted out faunal remains are scarce, gnaw marks are frequent and appear on almost all of the post-cranial bones

Figure 7 Examples of sawed, cut or chopped off (marked by arrows) reindeer antlers found melted out of Norwegian ice patches.

© Equinox Publishing Ltd. 2018 92 Jørgen Rosvold (e.g. Figures 5b, d and Figure 8). Rodent gnaw marks are rare, but many of the antler finds bear the characteristic marks of herbivore (likely reindeer) gnawing on the tines (e.g. Figure 8a). Herbivores are known to chew bones and antlers in order to extract valuable minerals (Wika 1982; Cáceres et al. 2011). Most gnaw marks, however, can easily be ascribed to large carnivores. Few crania show the characteristic tooth marks from carnivore gnawing, but the maxilla and nasal bones are usually lost, a typical feature on gnawed skulls (Haynes 1980), and most mandibles have been cracked open to extract marrow (Figures 5b and 8b). Thus far, we have found bones of arctic fox (Vulpes lagopus), brown bear (Ursus arctos) and wolverine (Gulo gulo) at alpine ice patches (Figure 3), but the information on how such sites are utilized by predators as potential hunting grounds or food caches is scarce (Rosvold 2016). Wolverines are known to use alpine ice to store carcasses especially during the summer (Bevanger 1992; Inman et al. 2012), and several of the finds have canine puncture marks that are consistent with wolverine teeth (e.g. Figures 5d and 8c). It thus seems highly likely that a large part of the faunal material are the melted out remains of wolverine food caches, but it is possible that other predators might have contributed as well.

Discussion Despite receiving much less attention than the purely archaeological finds, the amount, diversity and geographic distribution of glacial faunal finds are starting to reveal a rich source of ecological and historical data from around the world. The radiocarbon dates from the Scandinavian finds show that this record goes back at least 4700 years BP (Figure 4, Appendix 2), while results from the extensive Yukon material extends even further back in time to at least 9300 years BP (Hare et al. 2012). The use of this data, however, relies on a good understanding of its taphonomy, which should be studied in-depth at all sites. In order to understand both the archaeology and the wider ecology of the sites, it is especially important to know how the faunal material ended up on the ice. Do the finds reflect animals that actively spent time on the ice; i.e. animals which were part of the glacial ecosystem and who represented potential resources that attracted humans to the sites? Alternatively, do the finds represent animal remains that were brought to the ice through the actions of humans or other organisms, e.g. as food caches, offerings etc.? Or do the finds just represent accidental events of stray animals, unrelated to the ecology and normal use of the ice? A first step would be to determine if the finds are “cultural” (i.e. deposited as a result of human actions) or “natural” (i.e. deposited from animals). This can be hard to achieve for all finds, but some general patterns are apparent from the hitherto published finds and the Norwegian material presented above. While it is evident that humans have used the alpine ice extensively for various reasons, including hunting, the evidence for a direct connection between the melted out faunal material and human activities are rare. Among the Norwegian material only c. 1.2% of the finds show marks from butchery, which is in line with what has been observed in other large collections (Andrews, MacKay, and Andrew 2012; Hare et al. 2012; VanderHoek et al. 2012). The interpretation of glaciers and ice patches as archaeological sites is made difficult by the fact that the material is not usually ex-

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 93 cavated from the spot where it was deposited. The material is usually found on the surface after having been moved around due to the flow of the ice or by melting out and sliding down with meltwater. Fragments from the same depositional event could thus be spread far apart or get mingled with other unrelated finds (e.g. Figure 5e).

Figure 8 (Top) Gnaw marks on the brow tine of a melted out antler, likely chewed on by a reindeer. (Middle) Melted out cattle mandible with gnaw marks (marked by arrow). (Bottom) Canine puncture marks on both sides of a melted out reindeer scapula fits perfectly with the teeth of a modern wolverine skull. Photo: Å. Hojem/NTNU University Museum.

© Equinox Publishing Ltd. 2018 94 Jørgen Rosvold It is thus rarely possible to draw conclusions based on the context of the faunal material in relation to archaeological artefacts. However, with a continued effort to docu- ment all faunal finds from the sites, the potential to use body part representation to infer carcass processing and transportation will increase. From the materials collect thus far though, it seems that most of what was hunted by humans on the ice was transported away and not processed at the kill sites. It is also worth considering that humans could have brought some of the more exotic finds (i.e. animals that do not normally occur in high alpine areas) up to the sites. Thus far, there are no clear indications of this, apart from a few fish remains and domestic animals, e.g. horses and mules (see above). The Norwegian glacial faunal finds are in general comparable with what has been found on similar sites around the world, indicating that the evidence to date largely points towards that most of the faunal finds from glaciers and ice patches being naturally deposited. This natural deposition could have happened in several ways: the animals could have actively visited the sites and died there or left traces of themselves; prey remnants could actively have been brought up to the sites by predators; or animals could have ended up on the ice unintentionally through stray movements or when crossing mountains. Examples of all of these seem to have been found and it is important to differentiate between them in order to understand the ice as an ecosystem. Alpine and arctic animals visit and actively use inland ice for several different activities (Rosvold 2016) and such species, and seasonal visitors, are by far the most common among the Norwegian finds as well as worldwide (Appendix 1). Dung, strips of velvet, shed antlers etc. are good evidence that the animals have spent time on the ice. Even complete mummified carcasses of these species, with no signs of predation, are good indications of this and could be of animals that fell into crevices or died of other natural causes on the ice. If well sampled, their relative abundance in the collections can show the importance of glaciated sites for these animals and how much time they spend on them. The large amounts of shed reindeer antlers are good examples of this and direct evidence for the active presence of these animals through time, at different seasons of the year and potentially sex biased selection of areas. What is left on the sites will, however, be dependent on how the sites are exploited. Animals that use the sites frequently, but not for long periods of time, like predators and birds foraging on insects, might not leave as many clearly evident traces of their activities as herd-living ungulates that stay there for large parts of the day. Their remains could be more in the form of dung, urine, hair and feathers and other small remains that might require environmental DNA analysis to detect and identify. Ad- ditionally, predators might also leave more indirect traces of their history in the ice through their prey remains. Carnivore gnaw marks are frequent on ungulate bones found at glaciers and ice patches in Norway and North America. These finds could be discarded prey items from animals caught on the ice or they could be remains from food caches. Food caching, in order to preserve food for later, is a well-known behavior among a range of bird and mammal species (Smith and Reichman 1984), but at such alpine ice sites it has only been described for wolverines (Inman et al. 2012).

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 95 Active hunting and prey processing on glaciers and ice patches by both predatory birds and mammals are behaviors that should be further explored in modern populations. In much the same way as humans, it is possible that animals use the predictable occurrence of certain prey species on the ice, at different times of the day and year, to locate their prey. This is certainly the case for various birds that feed on insects that have become stuck on the ice (see Rosvold 2016 for a review). If gnaw marks and other signs of predation can be correctly identified to the species that caused them, this can be used to indicate the active use of the ice by predatory species that are not represented by their own bones. The prey items themselves cannot be used to infer the active presence of the prey species on glaciers or ice patches, but are nevertheless valuable data for alpine history in general. Among the Norwegian material, and the reported finds worldwide, there is a relatively large number of both alpine and non-alpine species of whom we are not aware of any behaviors linked to glaciers or ice patches (Appendix 1). Many birds make energetically costly mountain crossings on their seasonal migrations and die while migrating (Newton 2007). If they happen to fall down on perennial ice, they can be preserved for long periods of time. Such occurrences and random straying may represent a diverse array of non-alpine species. Finds of these species will likely be lim- ited to just one or a few individuals at each site, and this seems to be the case for most of the reported finds thus far. Random straying could also explain the many mummified small rodents, e.g. lemmings (Lemmus spp.) and voles (Microtus spp.), that have been found on the ice in both North America and Norway (Appendix 1, Figure 3). Alternatively, these more cryptic finds could indicate some hitherto unknown behaviors among certain species that will require more observational studies of glaciers and ice patches. In that regard, the ice finds can aid in defining species and areas where such behaviors should be studied in modern populations.

Conclusions and future prospects The view that glaciated landscapes are not barren, lifeless wastelands, but actually diverse and distinct ecosystems is slowly being recognized among researchers. Glacial archaeology has been and is important for including and understanding the role of humans and other vertebrates into these systems. Faunal finds from ice patch projects highlights species and behaviors tied to the ice for which we have few modern observations. The finds are starting to reveal that ice covered lands have been important areas for a wide range of vertebrates, including humans, for thousands of years. It is also clear that the ice represent an extraordinarily well-preserved record of faunal history for many species and material from such sites have already proven to have a significant information potential for genetic studies (Kuhn et al. 2010; Letts et al. 2012; Olivieri et al. 2014; Røed et al. 2014). In order to get better insight into the ecology of the ice itself, the diversity of use and deep-time significance to different organisms, it is important that all kinds of faunal remains are collected and reported, and not just the more spectacular finds. In order to fully characterize alpine ice as ecosystems we also need more com- parative studies between different kinds of sites. It is likely that various species, and

© Equinox Publishing Ltd. 2018 96 Jørgen Rosvold potentially sexes, value and use different types of ice covered landscapes based on differences in environmental factors like wind and moisture, and size, shape and sta- bility of the ice. For animals that use the ice to escape insects, larger ice patches might be preferred over small ones, or small ones might be selected if they are more exposed to wind. Various behaviors could also be tied to different areas on an ice patch. Ptarmigan (Lagopus spp.) are known to use cavities along the edge of ice patches to escape heat, but forage for windblown seeds and insects on the surface of the ice (Kaisila 1952; Johnson 1968; Pedersen et al. 2014). Knowledge about such differences would have been important for ancient human hunters as well. We also need to link vertebrates with the microorganisms in the ice to find out how and if they influence each other. Vertebrates transport valuable nutrients to the ice from outside areas and even the faunal remains themselves can be important parts of glacial ecosystems by leaking nutrients into the ice that may sustain microorganisms. It is even possible that certain microorganisms in the ice could negatively affect the health of visiting vertebrates (Edwards 2015). Alpine ice attracts a diversity of species from a wide area surrounding the sites (Rosvold 2016), while at the same time collecting and preserving traces of these species through time. By doing so, the ice serves as a sort of “baited pit trap” for natural history data on alpine organisms, and to a lesser degree as a source of data for non- alpine species. The fact that many of the animals use the sites actively makes the inland ice distinct from the classic paleontological pit traps, like sinkholes and tar pits, which trap animals in a more random manner. The type of material stored in the ice will also differ from these types of sites in that it not only “traps” complete organisms, but several kinds of remains from living organisms as well. As most of the material is naturally deposited, the faunal collection from the ice will be more biologically representative than what is found on archaeological sites. This, however, will depend on good collection strategies. In contrast to most archaeological finds, the material from glaciers and ice patches is not excavated and sifted. Instead, the material is found on the surface when walking on of the ice or along the edges. Small animal remains are harder to spot and care must be taken in order to get as representative a collection as possible. Many alpine ice sites contain both naturally deposited faunal material and cultur- ally deposited artefacts that are not necessarily directly linked. The sites themselves, are thus hybrids between natural historic and archaeological sites, and this duality has potential management implications as natural deposits often do not have the same protective management status as archaeological artefacts in many countries. If the faunal material is considered naturally deposited, it also stands the risk of losing the interest of many archaeological projects. This could potentially lead to less representative collection strategies for such finds unless the material receives increased attention among biologists or paleontologists. The material, however, offers unique contextual information about which potential resources were available to humans on the sites, at different times and in different areas. If some sites were more important for different kinds of animals at certain times of the year, this is likely reflected in various hunting strategies or other uses, like animal husbandry or food

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 97 storing, at different kinds of glacial sites or during various climates. Understanding the wider ecology of glaciated land is also essential for inferring the role of humans in these systems. Were more climatically stable sites more important to humans during warmer climatic periods when there were fewer alpine ice sites available and potentially greater densities of prey at the remaining sites? Were some sites used to harvest males vs. females; birds vs. ungulates? In order to do such comparisons it is essential to gather a representative collection of faunal finds from many sites. Faunal finds are often more abundant at glaciers and ice patches than archaeological artefacts (Andrews, MacKay, and Andrew 2012; Hare et al. 2012; pers. obs.) and detailed studies on the age distribution and preservation of faunal finds within sites will likely also provide relevant taphonomic information for archaeological finds. The alpine ice offers an interesting temporal duality that is paralleled in both the culture and nature of the ice. The ice is still an important agent in the lives of people today and serves as distinct and living ecosystems for many species. As the ice melts away it will affect a range of different organisms, including humans, but at the same time the inland ice is revealing its own history and deep-time importance. The faunal materials that melts out are vital in highlighting the fact that the ice itself is a habitat that offers a range of important ecosystem services. In order to understand the wider ecology of the alpine ice, we need to understand the history of both the culture and nature of the ice itself. This could even provide insights into Ice Age life, when even more of the landmasses were covered by ice. Climate change is both a blessing and a curse for such studies. The melted out remains offer a new and often uniquely well- preserved material that can offer paleoecological information unlike any other type of sites. However, if not rapidly collected the finds may quickly degrade and we risk losing a large amount of natural and cultural heritage with increased glacial melting.

Acknowledgements Financial support for the project was provided by the Norwegian Environment Agen- cy, The Research Council of Norway, the Sami Parliament, The Royal Norwegian So- ciety of Sciences and Letters, and Nansenfondene. Radiocarbon dates from south- central Norway was provided by Jostein Bergstøl at the Museum of Cultural History and I am very grateful to him, Per Å. Persson and Julian P. Martinsen, as well as Lars Pilø, at Oppland County Council, and Norsk Fjellmuseum for making these collections available. I am very grateful to all the local volunteer collectors, researchers and management institutions who have contributed with their finds and information, and especially Tord Bretten at the Norwegian Nature Inspectorate. Large thanks go to Greg Hare at the Government of Yukon for making the faunal finds from Yukon available, to Martin E. Callanan for valuable discussions and fieldwork, and to two anonymous reviewers for helpful feedback on the manuscript.

98 Jørgen Rosvold References 1 5 5 1 5 3 5 10 29 12 14 16 12 15

3,8 4,5 1-5 1,13

3,5-9 11,29 1,4,15

New Guinea New East Africa East

X X X Alps

X X X X X

Peru

Greenland Western U.S. Western

X X

America

NW North North NW X X X X X X X X X X X Scandinavia X X X X X Domestic cattle Unid. “antelope” Grey wolf Alpine ibex Wapiti Red deer African civet Domestic horse, mule North American porcupine Domestic cat Wolverine Norway lemming Siberian brown lemming N. A. brown lemming Snowshoe hare Mountain hare African wild dog Canada lynx Hoary marmot Common name Moose American bison Scandinavian finds and published reports of vertebrate specimen finds associated with glaciers and ice patches globally. Note that the taxonomy of some species have changed since original publications. New

Bos taurus Bovidae Canis lupus Capra ibex Cervus canadensis Cervus elaphus Civettictis civetta Equus spp. Erethizon dorsatum Felis silvestris Gulo gulo Lemmus lemmus Lemmus sibiricus Lemmus trimucronatus Lepus americanus Lepus timidus Lycaon pictus Lynx canadensis Marmota caligata Species Mammals Alces alces Bison bison Appendix 1

© Equinox Publishing Ltd. 2018 Faunal Finds from Alpine Ice 99 5 1 1 5 1 1 3 3 8 3 1 25 26 28 22 18 25 29 1,5 1,4 6-9 3-5 5,17 3,17 3,5,17 12,19,20 23,24,29 1,3-5,15,17,21,22 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Stephan’s dove Common raven Common quail Dall sheep Leopard Reindeer Alpine chamois Common shrew Arctic ground squirrel Bush duiker Brown bear Polar bear Red fox Arctic fox Hawk Razorbill Duck Plumed egret Field vole Long-tailed vole Tundra vole Northern red-backed vole Mule deer Muskrat Muskox Domestic sheep Bighorn sheep Eagle owl stephani

Corvus corax Coturnix coturnix Ovis dalli Panthera pardus Rangifer tarandus Rupicapra rupicapra Sorex araneus Spermophilus parryii Sylvicapra grimmia Ursus arctos Ursus maritimus Vulpes vulpes Vulpes lagopus Birds Accipitridae Alca torda Anserinae Ardea plumifera Microtus agrestis Chalcophaps Microtus longicaudus Microtus oeconomus Myodes rutilus Odocoileus hemionus Ondatra zibethicus Ovibos moschatus Ovis aries Ovis canadensis Bubo bubo

© Equinox Publishing Ltd. 2018 100 Jørgen Rosvold 5 1 1 1 1 1 1 2 1 25 27 25 25 25 25 25 25 12 25 1,5 2008. 23: Tyndall 1871. X X X X X X X X X 2007. 22: Knudsen et al. et al. X et al. 1996. 15: VanderHoek et al. 2012. 16: Döppes et al. 2014. 14: Weisgram 14: 5: Hare et al. 2012. et 6: Pattie al. 1967. 7: Lee et al. 2006. 8: Lee 2012. 9: Lee et al. X X X X X X X X X X X 3: Farnell et al. 2004. et 4: Andrews al. 2012. 2006. 12: Guest and Leedal 1954. 13: Savioz 2015. Savioz 13: 1954. Leedal and Guest 12: 2006. et al. Oriental cuckoo Papuan mountain-pigeon Moustached treeswift Dusky grouse Woodpecker White-winged diuca finch Oriental dollarbird Rock ptarmigan Ptarmigan Crossbill Little curlew Far Eastern curlew Red-necked phalarope Warbler Red-billed quelea Paradise kingfisher Western capercaillie Redwing Chum salmon Atlantic mackerel 2005. 18: and Bennike Andreasen 2005. 19: Latham 1926. 20: Mizuno 2005. 21: VanderHoek albertisii

mystacea

saturatus

1: This paper. 2: Beattie et 1: This paper. al. 2000. 2012. 10: Meyer 1891. 11: Lüps 17: Dixon et al. 24: Schlüchter et al. 2006. 25: Schodde et al. 1975. 1995. 26: Zbären 27: Hardy and 2008. Evans 1993. Naturmuseum 28: Young 29: Database of Bündner ID-nos. BNM 014010, 014567, 014635, 016820, 016828 and 016839. Dendragapus obscurus Dendrocopos sp. Diuca speculifera Eurystomus orientalis Lagopus muta Lagopus sp. Loxia sp. Numenius minutus Numenius madagascariensis Phalaropus lobatus Phylloscopus sp. Quelea quelea Tanysiptera spp. Tetrao urogallus Turdus iliacus Fish Oncorhynchus keta Scomber scombrus Cuculus Gymnophaps Hemiprocne

Radiocarbon Lab # SD Species Location date TRa-10216 1,012 0,001 Reindeer Storgrovfonna TRa-10189 1,049 0,002 Reindeer Fonnfjellet TRa-10240 1,06 0,002 Norway lemming Storbreen TRa-10217 1,064 0,001 Reindeer Storgrovfonna TRa-10260 1,069 0,002 Arvicolinae Kringsollfonna TRa-10235 1,069 0,001 Atlantic mackerel Østre Kringsoll TRa-10225 1,0824 0,002 Ptarmigan Kringsollfonna TRa-10203 1,094 0,002 Reindeer Storskardbreen TRa-10655 1,132 0,001 Reindeer Kringsollfonna TRa-10249 1,135 0,001 Razorbill Storbreen TRa-10262 1,199 0,003 Fur sample Sandskaret TRa-10202 1,261 0,002 Reindeer Storskardbreen Ua-50166 1,326 0,006 Reindeer Kringsollfonna TRa-3508 1,331 0,005 Reindeer Storbreen TRa-10245 1,34 0,003 Norway lemming Storbreen Ua-50148 1,41 0,006 Reindeer Storbreen TRa-10657 1,463 0,002 Reindeer Kringsollfonna TRa-10237 1,567 0,002 Arctic fox Knutshø Ua-50167 1,581 0,007 Reindeer Kringsollfonna TRa-10246 1,591 0,002 Norway lemming Storbreen TRa-3506 30 30 Reindeer Geithøtta Ua-50161 49 32 Reindeer Storbreen TRa-10252 70 25 Ptarmigan Kringsollfonna Ua-50163 79 32 Reindeer Storbreen TRa-10201 91 8 Reindeer Austre Krutfjellet Ø TRa-10197 92 17 Reindeer Kringsollfonna TRa-10205 95 13 Reindeer Storskardbreen TRa-10255 97 26 Norway lemming Kringsollfonna TRa-10219 120 30 Reindeer Evighetsfonna TRa-10204 127 7 Reindeer Austre Krutfjellet N TRa-10236 144 16 Cattle Knutshø TRa-10238 149 17 Norway lemming Storbreen TRa-10218 151 18 Reindeer Evighetsfonna TRa-10199 166 8 Reindeer Laupskardfjellet Ø TRa-10227 167 12 Rock ptarmigan Kringsollfonna TRa-10243 181 14 Sheep/goat Kringsollfonna Ua-44850 184 30 Reindeer Storbreen TRa-10239 191 13 Norway lemming Storbreen TRa-10231 195 19 Ptarmigan Kringsollfonna

TRa-10254 208 16 Rodent Storbreen TRa-10228 223 21 Rock ptarmigan Kringsollfonna TRa-10261 228 26 Reindeer Storskardbreen TRa-10233 237 14 Warbler Storbreen TRa-10248 277 12 Sheep/goat Kringsollfonna Ua-44852 335 30 Reindeer Storbreen Ua-50160 339 33 Reindeer Storbreen TRa-10214 356 15 Reindeer Blåhøa TRa-10232 357 14 Woodpecker Storbreen TRa-10241 363 14 Norway lemming Storbreen TRa-10230 374 13 Ptarmigan Kringsollfonna TRa-10242 388 18 Field vole Storbreen Ua-44848 396 30 Reindeer Knutshø TRa-10213 405 23 Reindeer Speilsalfonna TRa-10652 446 16 Reindeer Kringsollfonna Ua-50152 454 30 Reindeer Storbreen Ua-50144 456 32 Reindeer Storbreen Ua-50164 483 31 Reindeer Lertjønnkollen TRa-10656 484 13 Reindeer Kringsollfonna Ua-44854 505 30 Reindeer Lertjønnkollen TRa-10653 531 16 Reindeer Kringsollfonna TRa-10229 568 14 Rock ptarmigan Kringsollfonna TRa-10221 627 14 Reindeer Bjørnabotnhøgda Ua-50170 635 31 Reindeer Midtre Gjevilvasskam Ua-47116 650 35 Reindeer Lendbreen Ua-50162 673 34 Reindeer Storbreen Ua-50165 688 31 Reindeer Kringsollfonna TRa-10198 705 10 Reindeer Sanskardtjønna TRa-10191 724 13 Reindeer Kringsollfonna Ua-44851 825 30 Reindeer Storbreen Ua-47120 845 35 Reindeer Lendbreen TRa-10187 863 14 Reindeer Hoemsbreen Ua-47117 865 31 Reindeer Lendbreen Ua-50172 868 34 Reindeer Midtre Gjevilvasskam Ua-47115 870 35 Reindeer Lendbreen TRa-10253 934 20 Rodent Storbreen Ua-50173 936 33 Reindeer Midtre Gjevilvasskam TRa-10200 949 11 Reindeer Laupskardfjellet Ø Ua-50153 956 30 Reindeer Storbreen TRa-10184 1032 15 Reindeer Kringsollfonna TRa-10206 1040 21 Reindeer Grovåskardet TRa-3504 1065 30 Reindeer Storbreen TRa-10211 1100 12 Reindeer Grovåskardet TRa-10212 1103 21 Reindeer Grovåskardet TRa-10659 1107 14 Reindeer Kringsollfonna TRa-10207 1116 31 Reindeer Grovåskardet TRa-10210 1116 22 Reindeer Grovåskardet

TRa-10244 1121 18 Wolverine Lertjønnkollen Ua-44849 1128 30 Reindeer Knutshø TRa-10208 1173 30 Reindeer Grovåskardet TRa-10247 1185 17 Brown bear Storgrovfonna Ua-50169 1189 33 Reindeer Kringsollfonna TRa-10209 1222 17 Reindeer Grovåskardet TRa-10259 1230 26 Reindeer Løpesfonna TRa-10185 1245 17 Reindeer Evighetsfonna Ua-50171 1313 34 Reindeer Midtre Gjevilvasskam TRa-3507 1320 35 Reindeer Løpesfonna TRa-10224 1373 12 Duck Løpesfonna TRa-10196 1450 13 Reindeer Kringsollfonna Ua-47119 1465 33 Reindeer Lendbreen TRa-10250 1560 15 Common shrew Storbreen TRa-10188 1595 22 Reindeer Bjørnabotnhøgda TRa-10215 1621 36 Reindeer Blåhøa, Storfonna Ua-47113 1630 35 Reindeer Lendbreen Ua-50156 1672 30 Reindeer Storbreen TRa-10193 1721 13 Reindeer Kringsollfonna TRa-10192 1775 15 Reindeer Kringsollfonna Ua-47121 1820 38 Reindeer Lendbreen TRa-10223 1827 19 Reindeer Artfjället TRa-10664 1900 22 Reindeer Storbreen Ua-50145 1903 33 Reindeer Storbreen TRa-10654 1939 22 Reindeer Kringsollfonna TRa-10658 2005 15 Reindeer Kringsollfonna Ua-50155 2107 31 Reindeer Storbreen Ua-50146 2153 33 Reindeer Storbreen TRa-10660 2198 24 Reindeer Skirådalskollen Ua-44847 2221 30 Reindeer Stavsjølia Ua-47112 2235 40 Reindeer Åndfonne Ua-50168 2303 31 Reindeer Kringsollfonna Ua-50151 2448 30 Reindeer Storbreen Ua-47111 2455 35 Reindeer Åndfonne TRa-10195 2465 18 Reindeer Kringsollfonna TRa-10661 2475 19 Reindeer Storbreen TRa-10194 2478 13 Reindeer Kringsollfonna Ua-47110 2775 35 Reindeer Åndfonne TRa-10222 2704 10 Reindeer Midthøbreen Ua-50158 2891 38 Reindeer Storbreen TRa-3505 2915 40 Reindeer Brattfonna

Ua-50159 2922 36 Reindeer Storbreen Ua-47122 2928 48 Reindeer Lendbreen Ua-50147 2951 33 Reindeer Storbreen Ua-47114 3030 35 Reindeer Lendbreen TRa-10226 3064 21 Western capercaillie Storbreen TRa-10663 3101 20 Reindeer Storbreen TRa-10665 3101 21 Reindeer Storbreen TRa-10186 3123 15 Reindeer Kringsollfonna Ua-50150 3198 32 Reindeer Storbreen TRa-10183 3370 18 Reindeer Storbreen Ua-47118 3443 35 Reindeer Lendbreen Ua-50149 3484 40 Reindeer Storbreen Ua-50154 3539 34 Reindeer Storbreen TRa-10662 3541 26 Reindeer Storbreen TRa-10220 3870 18 Reindeer Evighetsfonna Ua-44846 3888 30 Reindeer Knutshø TRa-10234 4054 19 Western capercaillie Storbreen

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