Response distances of wild forest reindeer (Rangifer tarandus fennicus Lönnb.) and semi-domestic reindeer (R. t. tarandus L.) to direct provocation by a human on foot/snowshoes

Rangifer, 33, (1) 2013 32 (1), 2012 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no Response distances of wild forest reindeer (Rangifer tarandus fennicus Lönnb.) and semi-domestic reindeer (R. t. tarandus L.) to direct provocation by a human on foot/snowshoes Mauri Nieminen Finnish Game and Fisheries Research Institute, Reindeer Research Station, Toivoniementie 246, FIN-99910 Kaamanen (Corresponding author: [email protected]) Abstract: The objective of the study was to examine response distances of wild forest reindeer (Rangifer tarandus fennicus Lönnb.) and semi-domestic reindeer (R. t. tarandus L.) in Finland and Norway to direct provocation by a human on foot/snowshoes in 5 areas and in 15 reindeer herding cooperatives during different seasons in 2010-12. There were no significant differences in mean herd size or in sight, alert, flight and closest response distances of wild forest reindeer in the Kuhmo and Suomenselkä areas. The encounter distance in wild forest reindeer was significantly (P< 0.005) longer than in semi-domestic reindeer in Finland and in Finnmark, Norway, and it increased with the group size. The sight and the alert distances in wild forest reindeer were significantly (P< 0.001) longer than in semi-domestic reindeer. In addition, the flight distance for wild forest reindeer (mean 192 m) was significantly (P< 0.001) and almost three times longer than in semi-domestic reindeer in Finland (mean 68 m). The closest mean distance was in wild forest reindeer 191m (range 100-320 m) but only 44 m (range 2-110 m) in semi-domestic reindeer (P< 0.001). The sight, alert, flight and closest response distances were slightly longer in Norwegian than in Finnish semi-domestic reindeer. However, these distances were significantly (P<0.005) longer in Pohjois-Salla (no supplementary feeding) than in other Finnish reindeer herding cooperatives and at the Kaamanen experimental station. The mean flight distance of reindeer in Pohjois-Salla was 115 m but only 65 m in other cooperatives (P< 0.001). The closest distance of semi-domestic reindeer in PohjoisSalla (mean 105 m) was more than 2.5 times longer than in other reindeer herding cooperatives (mean 40 m). The mean sight, alert and flight distances in wild forest reindeer in autumn and winter were significantly longer (P<0.005) than in semi-domestic reindeer in Finland. However, during summer these distances in wild forest reindeer herds with young calves were significantly longer (P<0.005). The mean herd size of Finnish semi-domestic reindeer was almost the same in different seasons, but in wild forest reindeer it was slightly bigger during winter and spring and smaller during summer and autumn, only 7-23 reindeer. The mean encounter and sight distances in semi-domestic reindeer were significantly longer (P<0.005) in winter, but the mean alert and flight distances were almost the same in winter and summer and slightly longer than during other seasons. The results suggest that the supplementary feeding practice during winter may likely cause a reduction in flight distances in semi-domestic reindeer. Key words: human disturbance; response behaviour; flight distance; Rangifer tarandus; wild forest reindeer; semidomestic reindeer; seasons; supplementary feeding Rangifer, 33, (1), 2013: 1-15 1 Rangifer, 33, (1) 2013 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no 32 (1), 2012 Introduction There are many activities in which people may negatively influence the behaviour of wild Rangifer tarandus, and also affect their movement and subsequent range use (Wolfe et al., 2000). Human activities and infrastructure contribute with noise for example from power lines, generators, windmills, and also from moving objects like humans on foot, snowshoes and skis, snowmobiles, four-wheelers, cars, aircrafts and helicopters. While roads alone are not likely perceived as a threat to reindeer, increasing roads and traffic are. During the last 30-40 years outdoor ecotourism, hiking, skiing and hunting have been expanding and increasing activities in more remote areas, including mountain habitats of wild and semi-domestic reindeer (Helle & Särkelä, 1993; Colman et al., 2001; Reimers et al., 2006). There have also been many changes in reindeer herding and husbandry practices over the last years, especially in Finland (Nieminen, 2006). According to Baskin & Skogland (2001) reindeer are at an early phase of domestication, but semi-domestic reindeer generally exhibit more relaxed fright and flight behaviour compared to wild reindeer (Reimers et al., 2000; 2006). Therefore, when comparing behaviour between different reindeer herds, it is important to know the origin and history of the herds in question. While the caribou subspecies are wild, the Fennoscandian tundra reindeer population includes many domesticated herds (Reimers & Colman, 2006). The semi-domestic reindeer herds in Northern Finland, Sweden and Norway, as well as the wild reindeer herds (with a mix of wild and domesticated origin) in Southern Norway, are originally Eurasian wild tundra reindeer (Rangifer tarandus tarandus L.). DNA-analyses by Røed et al. (2008) support independent origin of semi-domestic reindeer in Fennoscandia and Russia. The domestic gene pools seem to meet only in eastern Finland, mainly in Halla reindeer herding cooperative. However, the wild forest reindeer (R. t. fennicus Lönnb.) population in eastern Finland and the wild reindeer populations in central Norway have contributed little or nothing to the domestic gene pool (Røed et al., 2008). Domestication is the first step of selection, and it is argued that domestication has mostly resulted in quantitative rather than qualitative changes (Mignon-Grasteau et al., 2005). Human activities affect reindeer/caribou through the senses of hearing, sight and smell. According to Flydal et al. (2001) the hearing capacity of reindeer ranges from 70 Hz to 38 kHz at a sound pressure of 60 dB. It means that almost all noises and vocalizations are readily perceived by reindeer. Reindeer as other ungulates has apparently also very good day and night vision. Reindeer most likely perceive colours, but no particular colour appears to be dominant, and reindeer probably are unable to distinguish between red and green colour. It is mainly contrasts and movements that betray human presence. Because the eyes of reindeer are laterally positioned, the combined visual fields of both eyes cover virtually 360o (Nieminen, 1994). It means that reindeer can also spot predators and humans sneaking up from behind. Reindeer´s laterally positioned eyes limit, however, the binocular visual field. Although reindeer´s sense of smell is not well documented, the capacity to capture scents even under unfavourable wind conditions is well known by reindeer/caribou hunters, hikers and also reindeer herders. Sometimes smell alone can trigger flight of reindeer without input from other senses. Reimers & Colman (2006) predicted that reindeer would also respond at greater distances to the directly approaching person when the wind carried the human scent to the reindeer than when the reindeer could not smell the human intruder. The strongly elevated nasals of the wild forest reindeer living only in forested regions indicate the very keen sense of smell due to increased olfactory mucous membranes (Nieminen, 1980). 2 Rangifer, 33, (1) 2013 32 (1), 2012 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no It has long been recognized that learning plays an important role in the manner and degree to which ungulates respond to humans (Geist, 1971), and there are usually three major learned responses which also are valid for reindeer: habituation, attraction and avoidance. Domestication, habituation and sensitisation are essential in shaping adaptability of reindeer and caribou (Reimers & Colman, 2006). Many impact studies focus on the behavioural responses of wildlife to humans, because these attributes are generally more amenable to study than other forms of response (Bejder et al., 2009). The cases of presumed habituation or sensitisation may actually represent differences in the tolerance level of wildlife to anthropogenic activity. For example reindeer show decreased flight responses in areas with relative high amounts of human activities (Colman et al., 2001; Reimers et al., 2009), indicating the ability to habituate to human activities. Habituation to humans would occur more readily in caribou populations that were not hunted and lived in areas lacking natural predators (Klein, 1980; Aastrup, 2000). Habituation and also former experiences with predators significantly influence an ungulate´s perception of threat. The populations with few predators flushed at greater distance than those where predators are common. All predator studies reviewed by Stankowich & Blumstein (2005) classified humans as the predator and measured differences in flight initiation distance between ungulate populations that differed with regard to human exposure. Wild ungulate populations like Svalbard reindeer (R. t. platyrhynchus Vrolik) exposed to a relatively high level of human activities have become habituated to humans in a non-threatening context (Tyler, 1991). They are likely to perceive less risk when approached by humans than would animals in populations where encounter with humans are rare (Colman et al., 2001; Lund, 2008). Even though semi-domestic reindeer are disturbed by human activities, they can increase their tolerance towards humans if insect harassment is severe during summer (Skarin et al., 2004). Rutting activities during autumn obviously also affect reindeer behaviour more than the directly approaching human observer (Reimers et al., 2006). Flight initiation distance is the distance at which an animal begins to flee from an approaching predator or human. It is usually used in studies, because it is easy to measure and correlates with other key aspects of escape behaviour e.g. alert distance (Blumstein et al., 2005). According to Vistnes & Nellemann (2008) accurate assessment of impacts from human activity requires, however, regional-scale and usually long-term studies. The objective of the present study was to examine response distances of wild forest reindeer and semi-domestic reindeer in Finland and Norway to direct provocation by humans on foot or on snowshoes. The observations were collected during different seasons in areas subjected to combinations of high or low human activity, supplementary winter feeding or freely-grazing in the forests or on the fields. The objective was also to determine if flight distances have changed over the last years in response to increased human activity and supplementary feeding of semi-domestic reindeer on natural pastures or in corrals. Material and methods Study areas and reindeer herds In total, 55 different reindeer herds were included in the present study: 17 wild forest reindeer herds (Rangifer tarandus fennicus Lönnb.) in the Suomenselkä area (9 in Perho, 1 in Kyyjärvi and 7 in Alajärvi municipalities) and 3 in Kuhmo municipality in the Kainuu area, 32 semi-domestic reindeer herds (R. t. tarandus L.) in 15 reindeer herding cooperatives in Finland including a herd at Kaamanen experimental station (150 reindeer, fenced area, 44 km2), and 3 Rangifer, 33, (1) 2013 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no 32 (1), 2012 3 herds in Northern Norway (2 in Kautokeino, West-Finnmark and 1 in Pykeija, East-Finnmark) (Fig.1). Approximate herd size was 40 animals in wild forest reindeer and 115 animals in semi-domestic reindeer in Finland and Norway. The herds of wild forest reindeer were found with the help of GPS-collared females. In the studied reindeer herding cooperatives in Finland total numbers of counted reindeer varied between 2200-8300 animals. Reindeer herds were studied during different seasons in the central areas of the cooperatives. According to the reindeer herders these reindeer herds represent a mixture of animals owned by many herders (the animals have different earmarks), i.e. constitute of animals belonging to the main herd of the respective cooperative. The wild forest reindeer disappeared from Finland for decades in the beginning of the 1900s. They came back to the Kuhmo area in the 1950s. During 1979-80 two forest reindeer males and eight females were transferred from Kuhmo to Salamajärvi National Park (Nieminen & Laitinen, 1983) and since then the population has increased in the Suomenselkä area to over 1100 individuals. However, dur4 Fig.1. Study areas of wild forest reindeer in Kuhmo (64o 7´ N, 29o 31´ E) and Suomenselkä (Perho, Kyyjärvi and Alajärvi) (63o63o 13´ N, 24o24o 25´ E) and of semi-domestic reindeer in reindeer herding cooperatives (1 = Paistunturi, 4 = Muddusjärvi, 7 = Ivalo, 8 = Hammastunturi, 9 = Sallivaara, 10 = Muotkatunturi, 14 = Kyrö, 18 = Oraniemi, 23 = PohjoisSalla, 24 = Salla, 25 = Hirvasniemi, 28 = Poikajärvi, 46 = Oivanki, 49 = Pudasjärvi and 57 = Halla) (64o-70o N) and at the Kaamanen experimental station (69o 6´ N, 27o 11´ E) in Finland and in Kautokeino (69o N, 23o 2´ E) and Pykeija (69o 58´ N, 29o 38´ E) in Finnmark, Norway. Rangifer, 33, (1) 2013 32 (1), 2012 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no ing the last years the number of wild reindeer has decreased due to predation. In Kuhmo municipality, near the Russian border, the population of wild forest reindeer has also decreased, and the total number of reindeer is today about 900 (Kojola et al., 2009). Northern Finland differs from the south mainly by the type of forest, but also by the elevated watershed areas in central and eastern Finland. Suomenselkä and Kainuu belong to the middle boreal vegetation zone (Ahti et al., 1968). The landscapes are dominated by mires, and they are located on watersheds. Both areas are diverse in altitudes, mountains, hills and valleys, barren areas with forests, bogs and lakes. The landscapes are dominated by Norwegian spruce (Picea abies) and Scots pine (Pinus sylvestris) forests with ericaceous heather, lichen (Cladonia, Cladina spp.) and boggy areas. The northern reindeer herding cooperatives are situated in the north boreal vegetation zone (Ahti et al., 1968). The terrain is dominated by rolling hills with different aged forest stands of mainly Scots pine. Mountain birch (Betula pubescens czerepanowii) grows in the slopes of the highest hills, and only the tops of the highest fells are barren. The cooperatives in the middle and southern parts of reindeer herding area belong, like Kainuu, to the middle boreal vegetation zone (Ahti et al., 1968). Agriculture is common and there are many fields in the reindeer herding cooperatives and also in wild forest reindeer areas. Because the numbers of wild forest reindeer have decreased as described above, no reindeer was hunted during the study period in the Kuhmo and Suomenselkä areas. In the reindeer husbandry area in Finland, and also in Finnmark, Norway, lichens pastures are generally strongly or very strongly worn (lichen biomass < 100-300 kg dry weight/ ha) (Kumpula et al., 2009; Mattila, 2006). As results of reindeer grazing, lichen ranges in mountain areas, large national parks, nature reserves and other wilderness areas are worn in Finland (Nieminen, 2010). In the northern cooperatives also the reindeer summer pastures are worn, and grazing has been the main reason to change vegetation and cause erosion in some places. The current condition of winter pastures in combination with a continuous state of change for the worse show that maintaining the current number of reindeer on their natural winter pasture is no longer possible. However, the body condition of reindeer in Finland has usually been good even during hard winters, due to the intensive supplementary feeding practised in the cooperatives. Totally over 40 million kg feed (calculated as dried hay) are used yearly during winters for feeding of semidomestic reindeer, mainly on natural pastures or in corrals in the middle or southern reindeer herding cooperatives (Nieminen, 2006). Supplementary winter feeding of reindeer in Finnmark area, Norway, is not common, but in Finland only Pohjois-Salla reindeer herding cooperative has herded reindeer on the natural pastures without supplementary feeding during winters. The wild forest reindeer are freelygrazing in Suomenselkä and Kuhmo, and both winter and summer pastures are in rather good condition. In Kuhmo the amount of lichen biomass (dry weight) has over the last years been seven times higher than in the nearby Halla reindeer herding cooperative (Mattila, 2004). The road network in wild forest reindeer areas and also in reindeer husbandry areas are well developed, and every year some wild forest reindeer and over 4000 semi-domestic reindeer die by traffic. Also, some adult wild forest reindeer and more than 4000 semi-domestic reindeer are killed yearly by big predators in Finland (Nieminen, 2012). Data collection In 2010-12, during the four sampling periods of September-November (autumn and rutting period), February-March (winter), April (spring) and July (summer), a single and same 5 Rangifer, 33, (1) 2013 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no 32 (1), 2012 person (the observer) on foot or on snowshoes (used in deep snow), dressed in dark clothing, disturbed wild forest and semi-domestic reindeer during daylight hours by directly approaching them. The observer used binoculars and camera (Nikon D80) to document behaviour and places and measured later (using 1 m steps) response distances between the reindeer and the observer and the resultant distances by the reindeer after taking flight. Upon location of a group (≥ 4 reindeer), 10 parameters were recorded: 1) sample month, 2) group size (small: < 20 animals, medium: 20-49 animals, and large: >50 animals), 3) group composition (females and calves, males), 4) dominant activity of the group when first sighted (lying or grazing, rutting, moving), 5) wind direction relative to the observer (no wind, upwind or downwind), 6) vegetation type (open field, marsh, forest), 7) topography of the surrounding area (level, mountain, lake ice and feeding place), 8) visibility/weather (sunny, cloudy, raining/ snowing), 9) snow depth and 10) temperature. When a group of reindeer was first sighted, the observer took pictures, and used the so called direct approach method: advancing directly towards the centre of the group at a constant speed (about 4 km/hour) with < 10 second stops, to take pictures and later measured the four additional response distances defined below. When reindeer are first disturbed they show signs of awareness and fright by raising their heads and tails, urinating and sometimes jumping (deVos, 1960; Horejsi, 1981). The initial flight is often followed by curiosity behaviour: the disturbed reindeer circles around the intruder to catch the scent. All measurements were made from the position of the directly approaching observer to the nearest reindeer. The wildlife response distance terminology and methodology recommended by Taylor & Knight (2003) with the modifications following Reimers et al. (2003; 2009) were used in this study: 1) Encounter distance (END): the distance used as the starting point of the disturbance. The reindeer first discovers the provoker by sight or scent, indicated by looking, standing, turning their head or pausing from eating in a manner visible to the observer. 2) Sight distance (SD): the distance between the observer and closest reindeer when reindeer in the group displayed an alerted behaviour directed at the observer. 3) Alert distance (AD): the distance at which the reindeer group displayed an increased alert response by grouping together or by individuals urinating with one hind leg extended outward at an exaggerated angle, while staring at the directly approaching observer (Fig. 2). 4) Flight distance (FD): the distance from the directly approaching observer to the group when the reindeer initially took flight. 5) Closest distance (CD): the distance from the directly approaching observer to the nearest animal if a group approached the observer immediately before final withdrawal. 6) Escape distance (ED): the shortest straightline distance from where the reindeer took flight in response to the observer to where the reindeer resumed grazing or bedded down. 7) Assessment time: the time elapsed from alert to flight initiation estimated from measured distances and assuming a constant observer speed of about 4 km/hour. Statistical analyses Initially, data of all areas and both wild forest reindeer and semi-domestic were pooled to analyse the effect of each factor on different distances separately. Correlation among dependent variables was tested with original (untransformed) data using Spearman rank correlation. The response variables (END, SD, AD, FD and CD) were first transformed into their natural logarithms prior to analysis. Sight, alert, flight and closest distances were analysed with Mixed Models Analysis. Marginal F-tests were 6 Rangifer, 33, (1) 2013 32 (1), 2012 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no used for the full models contained area, season, group size, wind direction relative to the observer, vegetation type and dominant activity of reindeer. To test for differences for provocation methods and to relate independent variables to reindeer responses, a mixed, stepwise analysis of ANOVA (analysis of variance) was used. x2 -test was used to assess the relative seasonal frequency in group-size classes. The statistical differences response distances in different groups were tested using t-test. Statistical tests were carried out by use of SPSS ver. 7.0 for Windows. The data were examined for statistical significance at P<0.05. Results During the four sampling periods in 2010-12 totally 55 independent reindeer groups were encountered and used in statistical analysis. A total of 2216 reindeer were observed. Of these, 739 were wild forest reindeer and 1477 semidomestic reindeer. Although 27 reindeer groups were encountered during autumn, these groups represented only 22 % of the groups in the large size class (> 50 animals) and 9.3 % of all of the reindeer sampled. During autumn, rutting season, most reindeer were in few large, mixed-sex groups. Totally 14 groups were encountered during winter, 50 % of the groups was in the large size class and represented 20 % of all studied reindeer. During spring and summer, 8 and 6 groups, respectively, were encountered, and only 1-2 groups were in the large size class. During summer, reindeer were distributed usually in smalland medium-sized female-calf Fig. 2. The alert distance (AD) of a wild forest reindeer group in the Suomenselkä area, in February 2010. The reindeer group displayed an increased alert response by grouping together or by individuals urinating with one hind leg extended outward at an exaggerated angle, while staring at the directly approaching observer. Photo Mauri Nieminen. 7 Rangifer, 33, (1) 2013 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no 32 (1), 2012 groups. Mean herd size of wild forest reindeer was 37, which was almost the same as for semidomestic reindeer in Finland (mean 39). The groups observed in Norway (winter only) were bigger (mean 178 animals). The encounter (END), sight (SD), alert (AD), flight (FD) and closest distance (CD) were positively correlated between areas, with a general decrease in the correlation coefficients from SD to CD (Table 1). FD increased with increasing encounter distance (END) (Table 2), as did also SD and AD, indicating that when the observer approached reindeer from farther away they responded at longer distances. No significant differences, depending on whether the approach was made on foot or on snowshoes in winter, were seen in any of the response distances. There were also no significant differences depending on area (Kuhmo and Suomenselkä) in mean herd size, END, SD, AD, FD or CD of wild forest reindeer. The Encounter distance (mean ± standard deviation) of wild forest reindeer (332 ± 24 m) was, however, significantly (P< 0.005) longer than that of semi-domestic reindeer in Finland and Norway (226 ± 13 and 250 ± 6 m, respectively) (Table 3). END increased with group size in both wild forest reindeer and semi-domestic reindeer. SD (253 ± 16 m) and AD (216 ± 13 m) in wild forest reindeer were significantly (P< 0.001) longer than in Finnish semi-domestic reindeer (144 ± 7 and 94 ± 6 m, respectively). The FD in 8 Table 1. Spearman rank correlation for the five dependent variables encounter (END), sight (SD), alert AD), flight (FD) and closest distance (CD). Encounter distance Sight distance Alert distance Flight distance r P r P r P r P Sight distance (SD) 0.843 0.001 Alert distance (AD) 0.736 0.001 0.895 0.001 Flight distance (FD) 0.699 0.001 0.867 0.001 0.960 0.001 Closest distance (CD) 0.592 0.001 0.787 0.001 0.905 0.001 0.922 0.001 Table 2. Linear mixed-effects model for predicting flight-initiation distances (In transformed) of groups of wild forest reindeer and semi-domestic reindeer disturbed by anapproaching observer on foot/snowshoes in different areas in Finland and Norway in 2010-12. Variable Estimate SE df t-value P-value Sight distance, Intercept 3.748 1.229 26 3.05 <0.005 Season (winter vs. autumn) -0.555 0.463 26 -1.19 0.246 (winter vs spring) -0.307 0.604 26 -0.51 0.612 (winter vs. summer) -0.766 0.891 26 -0.96 0.348 Group size (large vs. small) 0.353 0.279 26 1.27 0.217 (large vs.medium) 0.157 0.205 26 0.77 0.451 Wind (no wind vs. upwind) 0.099 0.166 26 0.59 0.558 (no wind vs. downwind) 0.001 0.209 26 0.01 0.995 Vegetation type (open vs. forest) -0.12 0.564 26 -0.21 0.833 (open vs. marsh) -0.199 0.522 26 -0.38 0.706 Activity (rutting vs. feeding) 0.206 0.322 26 0.64 0.528 (rutting vs. lying) 0.008 0.231 26 0.03 0.974 (rutting vs. grazing) 0.003 0.205 26 0.02 0.987 Encounter distace (ln m) 0.327 0.089 26 4.11 <0.005 Rangifer, 33, (1) 2013 32 (1), 2012 This journal is published under the terms of the Creative Commons Attribution 3.0 Unported License Editor in Chief: Birgitta Åhman, Technical Editor Eva Wiklund and Graphic Design: Bertil Larsson, www.rangifer.no wild forest reindeer (192 ± 14 m) was almost three times longer than in semi-domestic reindeer (68 ± 5 m) in Finland (P<0.001) (Fig. 3). The escape distance (ED) of reindeer was possible to measure only in the mountain areas, and the mean ED of semi-domestic reindeer in Muotkatunturi reindeer herding cooperative in Finland and also in Kautokeino, Norway was 360 m. The mean distance between alert and flight was 24 m in wild forest reindeer and almost the same, 26 m in semi-domestic reindeer in Finland. With an encounter speed about 4 km/ hour, these distances suggest that there was a separation of 22-24 seconds, on average, from when the reindeer groups became alert until they took flight. The mean closest distance was 191 m (range 100-320 m) in wild forest reindeer but only 44 m (range 2-110 m) in semi-domestic reindeer in Finland (P<0.001). The mean SD, AD, FD and CD in Norwegian semi-domestic reindeer herds were slightly, but not significantly longer than in Finnish semidomestic reindeer. However, the mean SD, AD, FD and CD of the semi-domestic reindeer in Pohjois-Salla (no supplementary feeding) were significantly (P<0.005) longer than in the other Finnish semi-domestic herds (Table 4). The mean FD of the semi-domestic reindeer in Pohjois-Salla was 115 m, but only 65 m in the other herds. The mean CD of semi-domestic 9 Table 3. Observed response distances of wild forest reindeer and semi-domestic reindeer in Finland and also of semidomestic reindeer in Finnmark, Norway, when provoked by an observer on foot/snowshoes in 2010-12 (data pooled across years). Wild forest reindeer (Kuh, Suo) Semi-domestic reindeer (Fin) Semi-domestic reindeer (Nor) mean range n mean range n mean range n Herd size 37 4-152 20 39 11-121 32 116 72-220 3 Encounter (END) 332 120-600 20 226 100-400 32 250 232-255 3 Sight (SD) 253 100-450 20 144 80-250 32 200 198-205 3 Alert (AD) 216 100-320 20 94 50-160 32 150 145-156 3 Flight (FD) 192 100-320 20 68 30-120 32 135 120-150 3 Closest (CD) 191 100-320 15 44 2-110 26 120 100-140 2 Fig. 3. Response distances (mean ± SD) of wild forest reindeer (WFR) and of semi-domestic reindeer in Finland (SDR-Fin) and in Norway (SDR-Nor) in groups disturbed by a directly approaching human on foot/snowshoes in 2010-12. Sample sizes are: END (encounter), SD (sight), AD (alert), and FD (flight distance), n = 20 for WFR, n = 32 for SDR-Fin and n = 3 for SDR-Nor, and CD (closest distance), n = 15 for WFR, n = 26 for SDR-Fin and n = 2 for SDR-Nor. reindeer in Pohjois-Salla was 105 m, which was more than 2.5 times longer than in the other herds (mean 40 m) (Table 4). The mean END, SD, AD and FD of wild forest reindeer were long in autumn and winter, and significantly longer (P<0.005) than in semi-domestic reindeer in Finland (Table 5, Fig. 4). However, during summer these distances of wild forest reindeer herds with young calves were significantly longer (P<0.005). The mean herd size of Finnish semi-domestic reindeer was almost the same during the four