Habitat use and preferences may be subject to spatial and temporal changes. However, long-term studies of species–habitat relationships are the exception. In the present research, long-term trends in habitat use by an alpine ungulate, the Tatra chamois Rupicapra rupicapra tatrica, were analyzed. We examined how environmental changes attributable to climate change, removal of sheep, and habituation to hikers, which took place over the last half-century have changed the spatial distribu- tion of animals. Data on the localities of groups sighted between 1957 and 2013 during autumnal population surveys were used to evaluate habitat associations: these were correlated with year, group size, population size, and climatic conditions. The results indicate that the Tatra chamois is tending, over the long term, to lower its altitude of occurrence, reduce its average distance to hik- ing trails, and stay less often on slopes with a southerly aspect. These trends are independent of group size, population size, and the weather conditions prevailing during observations, though not for altitude, where increases in air temperature are related to ﬁnding chamois at higher elevations. The proportion of alpine meadows and slope in the places used by chamois is correlated with population size, while the proportion of areas with trees and/or shrubs is correlated with group size and air temperature, though long-term changes were not evident for these variables. To the best of our knowledge, this work is the ﬁrst to document long-term trends in habitat use by ungulates. It shows that a species’ ecology is inﬂuenced by human-induced changes: abandonment of pastur- age, high-mountain tourism, and climate changes, which constitute the most probable reasons for this aspect of behavioral evolution in the Tatra chamois. Key words: habitat selection, long-term study, population ecology, protected area, ruminant. Habitat use and preferences depend on the needs of animals and the Factors affecting habitat use by herbivores include weather condi- advantages accruing from their occupation of a particular place tions (e.g., temperature and precipitation), temporal and spatial (Krebs 2009; Macandza et al. 2012). However, these uses and pref- changes in resource distribution, abundance and quality (due to erences depend, in turn, on accessibility of habitats, the distribution, vegetation phenology and vegetation dynamics), accessibility of pla- and quality of key resources within them (Lott 1990; Arau ´ jo et al. ces of rest and concealment, individual features (e.g., sex, age, and 2011; Robertson et al. 2015) and also inter- and intraspecific inter- condition), and types of behavior (e.g., neophobia and neophilia) actions (Richard et al. 2010; Kiszka et al. 2012; Beest et al. 2014). (Clobert et al. 2012; Brivio et al. 2016; Ofstad et al. 2016). V C The Author(s) (2018). Published by Oxford University Press. 1 This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact email@example.com Downloaded from https://academic.oup.com/cz/advance-article-abstract/doi/10.1093/cz/zoy035/4983040 by guest on 13 July 2018 2 Current Zoology, 2018, Vol. 0, No. 0 Other influences include the presence of predators (Sergio et al. Materials and Methods 2003; Lone et al. 2014) and intraspecific competition (Beest et al. Study area 2014, 2016). The Tatras are the only mountains in central and eastern Europe of Needs and advantages, and hence habitat use and preferences an alpine character. They cover an area of some 800 km , around are not, however, a constant feature and may change in both time 20% of which lies in Poland, and the remainder in Slovakia. The ele- and space (Iversen et al. 2014). Short-term changes in the choice and vation difference is 1,755 m, with the tallest peak, the Gerlach, use of habitats are attributable mainly to the various facets of the 2,655 m amsl. The Tatras are protected in their entirety in the form growing season (Fynn et al. 2014; Thompson et al. 2014) or depend of national parks: the TatranskyNa ´ rodn y Park (TANAP) in on the reproductive cycle of herbivores (Nicholson et al. 1997; Slovakia (formed in 1949) and the Tatrzanski Narodowy Park (tatra Rolandsen et al. 2017). In contrast, long-term changes in habitat se- national park - TNP) in Poland (formed in 1954). In addition, the lection and use are associated with an animal’s life cycle and individ- Tatras have been declared a Man and Biosphere Reserve and are ual needs, which are governed mainly by changes in condition, body included in the Natura 2000 network of protected areas in Europe. size, the ability to reproduce, experience (Forchhammer et al. 2001; The Tatras are characterized by a vertical zonation of climate Long et al. 2016), or may be the result of its adaptation to changing and vegetation. The tree line lies at 1,500 m; above this, there are habitat conditions (Lister 2004; Kawecki 2008). alpine meadows and the subnival zone. This altitude corresponds to Most studies to date on changes in habitat preferences and use the cool, moderately cold, and cold climatic zones, with features have been carried out in the short-term context, chiefly in terms of including low air temperatures and low atmospheric pressure (Hess the seasonality of habitat conditions (winter–summer) or changes in 1996). The mean annual temperature is –0.7 C, and the coldest age cohorts (juveniles versus adults) over a small number of consecu- month is February. On average there are 188 winter days, that is, tive years (Brambilla et al. 2006; Nesti et al. 2010), exceptionally when the mean daily temperature (T )is <0 C, whereas a ther- mean decades (Garel et al. 2007; Shenbrot et al. 2010). A review of papers mal summer (when T >15 C) does not occur at all. The growing mean on habitat-related research from top-ranked ecology journals for the season (when T >5 C) lasts for just under 100 days. Snow typic- mean past 10 years shows that of 84 articles dealing with this subject, only ally covers the ground from December to April. In the alpine zone of 15% provide data for periods longer than 7 years, and barely 5% the mountains snow persists well into May and June, for an average concern long-term analyses of the nature of such changes (Uboni of 221 days of snow cover in the year, hindering access to grazing et al. 2015). There are a number of reasons why so few long-term areas and exacerbating the risk of triggering avalanches. In the high- studies have been carried out, even though they are fundamental to est elevation regions, the snow layer is no more than 50 cm thick understanding the ways in which animals adapt to their natural en- only from mid-June to early September; on occasion, snow falls in vironment: such studies require long-term commitments from scien- July and August. The mean annual precipitation is nearly 1,800 mm. tific institutions, permanent funding, and above all, untrammelled The thermal conditions prevailing in the Tatras, expressed as air access to study areas. This final criterion often means having to es- temperature, resemble those of the Alps (Nied zwied z 2006). tablish an uneasy working relationship with the owner or adminis- The Tatras are built of granitoids, metamorphic, and limestone trator of the land in question (Schradin and Hayes 2017). rocks. The rock formations in the zone inhabited by chamois can be The choice and use of optimal habitat present a challenge to divided into non-calcareous ones, from which acidic soils are montane ungulates living in extreme conditions. The occurrence and formed, and calcareous ones, which give rise to soils with a neutral survival of herbivore populations is determined by the availability of pH (Komornicki and Skiba 1985). The habitats in the zone occupied natural resources, which are strongly influenced by dynamically by chamois are associated with the plants Oreochloa disticha, changing climatic factors like precipitation and snow cover, often Juncus trifidus, Festuca versicolor, and Sesleria tatrae (Jamrozy varying widely from season to season (Shackleton and Bunnell et al. 2007). Above the tree line there is dwarf pine Pinus mugo 1987). The present analysis of the long-term changes in habitat use scrub: this is quite dense at lower altitudes, but gradually thins out by a montane ungulate is based on a long-term data set for Tatra with increasing elevation, becoming patchy and forming clumps, be- chamois Rupicapra rupicapra tatrica. During the last 50 years this fore disappearing entirely at around 1,800 m. As a result of the animal has been subjected to intense pressure owing to mass tourism abandonment of pasturage there has been regeneration of the wood- (Blazejczyk 2002; Skawin ski 2010): the spatial distribution of its land communities on mid-forest meadows, which lie at lower alti- population may have been affected by this. At the same time, protec- tudes, far below the range of occurrence of the Tatra chamois. Only tion of the Tatra Mountains in the form of a National Park has led locally, forest regeneration was recorded near the tree line and in the to the gradual elimination of pasturage from this area; it also guar- dwarf pine zone (Czajka et al. 2012). However, comparison of aer- antees the complete protection of the chamois population. Climate ial photographs from 1955 and 2004 shows that the course of the change (M. Ciach and Ł. Pe ˛ ksa, submitted for publication), as well tree line in the Tatras has not changed over the long term (Guzik as the dynamic changes in the size of chamois population — a dra- 2008) and no major changes in vegetation structure of the alpine matic fall from the 1950s until the turn of the century followed by the subsequent rebuilding of the population (Jamrozy and Pe ˛ ksa zone have been recorded. 2004) — could have affected habitat use as well. This paper analyses The zone occupied by chamois in the Tatras covers the area long-term changes in habitat use by the Tatra chamois and attempts above the tree line and reaches up to the summits of the highest peaks (Jamrozy et al. 2007). The areas above the tree line, lying in to identify the factors governing this. To do so, a unique and excep- their entirety within the two National Parks, are protected: hunting tionally long (57 years) series of data is used to track the changes in there is strictly prohibited and incidents of poaching are exceptional habitat use by these animals, providing an opportunity to assess the (Jamrozy et al. 2007). The Tatra chamois population is completely direction of changes in the ecology of the Tatra chamois. As far as we are aware, this material is one of the longest sequences of data isolated from other mountain areas, so neither emigration nor immi- for evaluating trends in habitat use by ungulates (see Festa-Bianchet gration are possible. A feature unique to the Tatra chamois popula- et al. 2017). tion is the lack of vertical migrations meaning that the animals Downloaded from https://academic.oup.com/cz/advance-article-abstract/doi/10.1093/cz/zoy035/4983040 by guest on 13 July 2018 Ciach and Pe ˛ ksa Long-term changes in habitat use 3 spend the whole year in habitats above the tree line (Jamrozy et al. were established for each chamois group location: altitude (accurate 2007). Within its distribution range, moreover, the Tatra chamois to 1 m), aspect (accurate to 1 ), and the slope of the terrain. The has no serious competition from other ungulates with regard to habitat variables were established for a circle of radius 100 m habitat and food resource use. The red deer Cervus elaphus and roe around the observation location. This radius was chosen because of deer Capreolus capreolus inhabiting the Tatras graze in the open the difficulty in defining an observation place as a point, because the (alpine) areas above the tree line only rarely and/or periodically chamois would often be on the move as they were being observed (Jamrozy et al. 2007). Sheep grazing, once common all over the (including while they were grazing), and in the case of larger groups Tatras, gradually declined following the creation of the National because of the scatter of the animals within. To calculate the value Park, and since the 1980s has been restricted to mid-forest mead- of a given variable the weighted mean was used by multiplying the ows, which lie at lower altitudes, far below the range of occurrence value of a pixel by their number in the circle. The slope layers and of the Tatra chamois (Mirek 1996). aspects were generated using functions available in the expanded Spatial Analyst (ESRI 2005). Aspect was expressed in eight classes (N, NE, E, SE, S, SW, W, and NW). For this analysis a variable, cal- Material and data sources culated as the summed area of pixels with S, SE, and SW aspects, The changes in habitat use by Tatra chamois were analyzed on the was used. The assumption was that slopes with a southerly aspect basis of counts done by the Tatra National Park (TNP) between usually have a longer growing season, thus providing the chamois 1957 and 2013. The counts were carried out in November (excep- with richer and more attractive grazing in autumn, and also that the tionally in early December, if the weather conditions so dictated). autumnal snow cover would be thinner and lie on the ground for a The counting methodology was based on the one suggested by Jozef shorter period, not hindering access to food. Mu ¨ ller in 1932 (Chudı ´k 1969). The entire area of the Tatras inhab- A vegetation cover map for the areas lying above the tree line ited by chamois was divided into counting areas that included all po- was also used. It was generated by means of an analysis of a satellite tential chamois habitats. Originally, the TNP was divided into 17 image taken by the Ikonos satellite in August 2004, and the areas such areas (14 in the High Tatras and 2 in the Western Tatras). above the tree line were allocated to 20 habitat classes (Guzik Later, up to 1988, their number was gradually increased to 30, their 2008). Variables were used to define the overall proportions of areas being reduced at the same time. In the history of these counts, alpine meadows (areas covered by grasses, sedges, herbaceous vege- there were years when none were undertaken (1979, 1981, and tation) and the overall proportion of shrubby and/or wooded habi- 1997) or the results (record cards), for some unknown reason, not tats (patches of dwarf pine, single trees, or clumps of trees growing deposited in the TNP archives (1958 and 2000). However, the main among dwarf pines or in open terrain). assumptions of inventories remain unchanged since their introduc- Distance to the nearest hiking trail was defined as the distance tion (Chovancova ´ et al. 2006) and the official Tatra chamois popu- between the position of the center of a group and the nearest trail. lation size in the TNP is based on the data acquired from these For this, the hiking trail layer (a total length of 275 km) in the annual inventories. possession of the TNP was used. The network of hiking trails in the Each TNP counting area was patrolled during two days by a Tatras was mainly determined in the period preceding the establish- team of at least two people, who recorded the numbers of chamois ment of the national park. The course of the trails, with minor in the groups encountered and the places where they had been modifications and periodic exclusion from use of selected trails, observed (Chovancova ´ et al. 2006). The results were recorded on remained relatively stable during the entire research period. cards, subsequently deposited in the TNP archives. Due to the legal Information on mean air temperatures, total precipitation, and restrictions (high regime of protection) and the conservation needs snow cover thickness in November 1957–2013 was provided by the (low population size), the animals remained unmarked. However, Institute of Meteorology and Water Management’s high-mountain the groups recorded by individual observers in a given counting sea- observatory at the summit of the Kasprowy Wierch mountain (alti- son were checked by the coordinator in order to prevent the multi- tude 1,987 m amsl), situated in the central part of the Tatra massif, plication of records of the same groups moving about and noted by in the center of the altitudinal zone inhabited by chamois. Overall different recorders. Based on the sex, age, number, and direction of population size of Tatra chamois was based on the data acquired movement multiplied records referring to the same group were from annual inventories (Chudı ´k 1969, see above for details) and excluded. During the count period the recorders, the coordinators represent official population estimates provided by national park summarizing the annual counts, and technical details of recording authorities. It is based on total sum of all individuals recorded dur- changed. For the purposes of this analysis, therefore, only those ing inventory (excluding the groups recorded multiple times). In records of chamois were used (N ¼ 2,425) that enabled the precise favorable conditions (rain-free and fog-free weather and cloud-base localization of groups (data include location on map, coordinates, height above summits) and with a large number of observers densely or precise description of the location) and that did not refer to the covering surveyed area this method assumes over 90% of the total same groups recorded multiple times. population to be counted (Chudı ´k 1969). However, the true detec- The data on animal observations were digitized to produce a tion probability of the method applied and, therefore, its accuracy layer with the autumn positions of the chamois and an attribute have not been tested at the time of its implementation and through- describing group size. Six habitat parameters were measured for out the entire monitoring period (Chovancova ´ et al. 2006). Since, each location where chamois were recorded: altitude above mean the number of animals in a given year could be underestimated sea level (m amsl), percentage of terrain with a southerly aspect (%), when reported based on counting, these estimates should be consid- slope (degrees), percentage of alpine meadows (%), percentage of ered to represent the minimum population size. habitats with trees and/or shrubs (%), and distance to the nearest hiking trail (m). The spatial analysis was carried out in ArcINFO software from the ArcGIS package (ESRI 2005) using the Data analysis 20 20 cm resolution numerical terrain model (NTM) in the posses- Descriptive statistics (mean6 SD, median with quartiles, and range sion of the TNP. On the basis of NTM, the following parameters values) of all six habitat variables — altitude above mean sea level, Downloaded from https://academic.oup.com/cz/advance-article-abstract/doi/10.1093/cz/zoy035/4983040 by guest on 13 July 2018 4 Current Zoology, 2018, Vol. 0, No. 0 Table 1. Descriptive statistics for all habitat variables analyzed at the Tatra chamois Rupicapra rupicapra tatrica observation sites and group size characteristics (TNP; for parameters, see the “Materials and Methods” section; N ¼ 2,425) Variable Mean SD Median Quartile range Range Altitude (m amsl) 1,899.3 180.0 1,930.9 1,796.9–2,034.3 1,294.7–2,332.1 Southerly aspect (%) 27.5 25.9 20.9 6.7–39.7 0.0–100.0 Slope (degrees) 38.3 8.4 39.0 33.4–44.1 12.7–61.2 Meadows (%) 37.5 31.3 28.4 9.7–64.0 0.0–100.0 Trees and shrubs (%) 6.3 16.1 0.0 0.0–1.3 0.0–98.2 Distance to trail (m) 293.1 292.0 185.2 43.3–485.4 0.1–1,001.0 Group size (individuals) 3.6 3.6 2.0 1–5 1–50 percentage of terrain with a southerly aspect, slope, percentage of There were significant long-term changes in habitat alpine meadows, percentage of habitats with trees and/or shrubs, parameters at the autumn observation sites with respect to the following variables: altitude (F ¼ 21.44, P¼ 0.000), southerly and distance to the nearest hiking trail — recorded in the places of 1,2417 aspect (F ¼ 17.85, P¼ 0.000), percentage of alpine meadows observation of Tatra chamois were calculated. In the first approach 1,2417 (F ¼ 4.48, P¼ 0.034), and distance to hiking trails (group level), the habitat parameters for the chamois record local- 1,2417 (F ¼ 22.08, P¼ 0.000) (Table 2 and Figure 1). Group size was ities (N ¼ 2,425) were taken to be dependent variables, and observa- 1,2417 unrelated to any of the habitat parameters except the habitats with tion year and group size were explanatory variables. Multiple shrubby and/or woody vegetation (F ¼ 4.18, P¼ 0.041) — regression analysis was used to evaluate the correlation between 1,2417 groups tended to be larger in areas with a larger proportion of habitat year, group size, and each of six habitat variables. containing patches of dwarf pine and/or clumps of trees (Table 2). The In the second approach (yearly means level), within-year mean effect size of statistically significant explanatory variables was small, values of each habitat variable were calculated (N¼ 52; excluding indicating a high level of variation in habitat use among Tatra chamois 5 years with missing data, see the “Material and data sources” sec- groups (Table 2). tion) and considered to be a new set of dependent variables. This ap- The long-term declining trend in the mean altitude of chamois proach was associated with the kind of explanatory variable used in observations (F ¼ 10.48, P ¼ 0.002) was counterbalanced by 1,46 the analysis: their values characterized the whole study area and the weather conditions at the time of observations (F ¼ 9.63, 1,46 were not attributed to any particular group. The observation year, P ¼ 0.003): higher temperatures in November meant that chamois the overall size of the chamois population estimated in a given year, could be seen at higher altitudes (Table 3). November temperatures and the weather conditions in November (the month when counting were also significantly correlated with the percentage of shrubby took place) in a given year, that is, temperature, precipitation, and and/or woody vegetation (F ¼ 6.87, P ¼ 0.012): as temperatures 1,46 thickness of snow cover, were treated as explanatory variables. fell, the chamois’ use of these habitats rose (Table 3). The chamois Multiple regression analysis was used to evaluate the correlation be- population size in the TNP and weather conditions in November tween these explanatory variables and mean values of each of the were not significantly correlated with the proportion of terrain six habitat variables. with a southerly aspect or with distance from hiking trails. This The strength of each predictor relying on effect sizes was esti- was indicative of the diminishing use of areas with a southerly mated using z-transformed Pearson’s product-moment correlation aspect (F ¼ 9.09, P ¼ 0.004) and closeness to hiking trails 1,46 coefficients. To describe the effect sizes of statistically significant ex- (F ¼ 14.66, P ¼ 0.000) over the long term (Table 3). Slope 1,46 planatory variables, the criteria listed by Cohen (1988) for small (F ¼ 17.91, P ¼ 0.000) and the proportion of alpine meadows 1,46 (r ¼ 0.1, explaining 1% of the variance), intermediate (r ¼ 0.3, (F ¼ 4.73, P ¼ 0.035) at the chamois observation sites depended 1,46 explaining 9% of the variance), or large effect sizes (r ¼ 0.5, explain- on population size: when the total number of individuals in TNP ing 25% of the variance) were adopted. To visualize long-term was larger, chamois were more likely to use areas with larger pro- trends in habitat use, least squares regression lines were fitted to portions of alpine meadows and gentler slopes (Table 3). The effect within-year means of each habitat variable with a significant trend. size of the statistically significant explanatory variables was inter- The statistical procedures were performed using Statistica 12.0 soft- mediate or large, indicating that they explain a considerable propor- ware (StatSoft Inc. 2014). All tests were considered significant with tion of the variance in inter-annual variation documented (Table 3). P< 0.05. Discussion Results This analysis of a 57-year-long series of data derived from autumn The mean altitude at which chamois were recorded in the TNP from counts of chamois shows that habitat use by these animals has 1957 to 2013 was 1,8996 180.0 m amsl (Table 1). The mean per- changed over the long term. The results indicate that chamois tend centage of terrain with a southerly aspect in the areas where the au- to be found at lower altitudes than they formerly did, they are seen tumnal observations were carried out was 27.56 25.9%, and these closer to hiking trails and do not use slopes with a southerly aspect localities had a mean slope of 38.36 8.4 . The mean percentage of as often as before. These trends are independent of group size, alpine meadows in these localities was 37.56 31.3%, and the mean population size, and weather conditions during the observation percentage of terrain with shrubby and/or woody vegetation was period (except with respect to altitude, where higher temperatures in 6.36 16.1%. The mean distance between the points of observation November were associated with an increase in the altitude at which of groups and hiking trails was 293.16 292.0 m (Table 1). chamois were observed). Downloaded from https://academic.oup.com/cz/advance-article-abstract/doi/10.1093/cz/zoy035/4983040 by guest on 13 July 2018 Ciach and Pe ˛ ksa Long-term changes in habitat use 5 2200 80 2 2 y = 4507.13 - 1.32x; r =0.14 y = 273.55 - 0.12x; r =0.16 1500 0 1957 1965 1973 1981 1989 1997 2005 2013 1957 1965 1973 1981 1989 1997 2005 2013 Year Year 54 90 52 y = -124.57 + 0.08x; r =0.04 28 0 1957 1965 1973 1981 1989 1997 2005 2013 1957 1965 1973 1981 1989 1997 2005 2013 Year Year 50 900 y = 4447.57 - 2.09x; r =0.23 0 0 1957 1965 1973 1981 1989 1997 2005 2013 1957 1965 1973 1981 1989 1997 2005 2013 Year Year Figure 1. Long-term changes in habitat use by the Tatra chamois Rupicapra rupicapra tatrica (TNP; for parameters, see the “Materials and Methods” section; N ¼ 2,425); dots and whiskers represent means and standard errors, respectively; and regression lines are shown for variables with a signiﬁcant trend (P< 0.05; see Table 2). The greater number of chamois records at lower altitudes is distribution to the presence of the sheep and the accompanying probably due to the gradual abandonment of the large-scale sheep sheepdogs (Chirichella et al. 2013). This study has also shown that pasturage in the areas where chamois are found (Mirek 1996). the long-term reduction in the altitude of chamois observation local- Competition with sheep is thought to be the main factor limiting the ities was moderated by the temperatures prevailing during the obser- Tatra chamois’ living space in lower-altitude alpine meadows vation period: in warmer years the chamois moved to higher (Jamrozy et al. 2007). A study in the Alps has shown that chamois elevations. Such movements in response to weather conditions are avoid places where sheep graze, instead adapting their spatial typical of animals in Arctic and high-mountain environments: their Downloaded from https://academic.oup.com/cz/advance-article-abstract/doi/10.1093/cz/zoy035/4983040 by guest on 13 July 2018 Shrubs and trees [%] Slope [degrees] Altitude [m] Distance to trail [m] Meadows [%] Southern aspect [%] 6 Current Zoology, 2018, Vol. 0, No. 0 Table 2. The results of multiple regression analysis testing for long-term changes in habitat use by the Tatra chamois R. rupicapra tatrica: correlation between year, group size, and habitat variables recorded at the group sighting location (TNP; for parameters, see the “Materials and Methods” section; N ¼ 2,425) Dependent variable Effect Estimate SE 95% CL þ95% CL tP Effect size Altitude Intercept 3,996.45 452.76 3,108.62 4,884.28 8.83 0.000 Year 21.06 0.23 21.50 20.61 24.63 0.000 0.094 Group size 0.08 1.00 21.89 2.05 0.08 0.936 0.002 Southerly aspect Intercept 305.55 65.59 176.93 434.18 4.66 0.000 Year 20.14 0.03 20.20 20.07 24.22 0.000 0.087 Group size 20.24 0.15 20.53 0.04 21.67 0.095 0.037 Slope Intercept 63.87 21.16 22.37 105.37 3.02 0.003 Year 20.01 0.01 20.03 0.01 21.19 0.234 0.026 Group size 20.08 0.05 20.17 0.01 21.71 0.087 0.036 Meadows Intercept 2129.70 79.29 2285.18 25.77 21.64 0.102 Year 0.08 0.04 0.01 0.16 2.12 0.034 0.042 Group size 20.18 0.18 20.52 0.17 21.01 0.315 0.019 Trees and shrubs Intercept 22.10 40.93 282.36 78.16 20.05 0.959 Year 0.00 0.02 20.04 0.04 0.19 0.851 0.005 Group size 0.19 0.09 0.01 0.36 2.04 0.041 0.042 Distance to trail Intercept 3,763.59 736.48 2,319.39 5,207.80 5.11 0.000 Year 21.74 0.37 22.47 21.02 24.70 0.000 0.097 Group size 22.65 1.63 25.85 0.55 21.62 0.105 0.036 Notes: Effect size is the z-transformed Pearson product-moment correlation coefﬁcient. Statistically signiﬁcant terms (P< 0.05) are shown in bold. intolerance of higher temperatures forces them to migrate to cooler within 1 km of a hiking trail (Blazejczyk 2002; Skawin ski 2010). areas (Aublet et al. 2009). In the case of the Tatra chamois, this The intensity of human pressure is exceptionally great around the implies movements toward the cooler alpine zone. The influence of Kasprowy Wierch mountain. In the peak summer season, the cable high temperatures on chamois migrations to higher (cooler) moun- car carries 1200–1300 people up to the summit daily, and a further tain regions is, however, regarded as relatively insignificant com- 2000–3000 people arrive there on foot (Pe ˛ ksa and Ciach 2015). pared with movements that are the consequence of flushing by sheep Such crowds of people are extremely stressful for the chamois, lead- and shepherds (Mason et al. 2014). ing to reactions measurable at both the physiological (elevated stress This study demonstrates a long-term decline in the proportion of hormone levels; Zwijacz-Kozica et al. 2013) and behavioral levels areas with a southerly aspect where chamois were counted in the au- (break-up of large chamois groups into smaller ones as a result of tumn. Areas with such an aspect have a milder climate and thus a their being flushed; Jamrozy et al. 2007). longer growing season (Toftegaard et al. 2016), which ensures more This study reveals that slope and the proportion of alpine mead- abundant food resources for a longer period (Seydack et al. 2012). ows at the chamois recording locations are correlated with popula- However, the global rise in average temperature (IPCC 1996)is tion size: the higher the number of animals, the more often they are gradually prolonging the growing season (Kullman 2004), as a result observed in habitats with a greater proportion of alpine meadows of which, areas with a southerly aspect are losing their advantage as and on terrain with a gentler slope. Chamois move onto steep slopes grazing places in autumn, since growing seasons on slopes with in order to minimize attacks by predators, which move far more other aspects are also getting longer. slowly on steep slopes than on flatter ground (Fox and Krausman Habitat use by Tatra chamois may be limited by extensive 1994). Utilizing steep slopes at times when overall numbers of Tatra human pressure (Pe ˛ ksa and Ciach 2015). At present the TNP is vis- chamois are low can thus reduce the effectiveness of predatory ited by 3 million people every year; as a consequence, the chamois attacks. When numbers are high, on the other hand, individual living there have had to adapt to the presence of human beings. The chamois groups are larger, and the animals use open habitats with a flight distance of the Tatra chamois from humans on hiking trails is gentler slope more often (Hebblewhite and Pletscher 2002): presum- less than 100 m (Jamrozy and Pe ˛ ksa 2004), which is less than that of ably owing to the dilution effect, but also because heightened vigi- the Alpine chamois Rupicapra r. rupicapra, for which the flight dis- lance lowers the hunting success of the predator (Lima and Dill tance was found to vary from 103 to 180 m (Gander and Ingold 1990). The changes in habitat use with increasing population size 1997). Our long-term data confirm that the distance separating could also be related to increasing intraspecific competition. Tatra chamois and hiking trails has systematically fallen over the The results of this research indicate that when temperatures are last half-century, indicating a gradually increasing tolerance of the low, chamois are more often found in habitats with larger propor- almost constant presence of large numbers of people on the Tatra tions of trees and/or shrubs. This may be because this taller vegeta- hiking trails. Even though the influence of hikers is largely restricted tion offers better shelter from inclement weather, especially strong to the hiking trails, it is worth noting that 96% of the TNP lies winds. However, these greater proportions of areas with trees and/ Downloaded from https://academic.oup.com/cz/advance-article-abstract/doi/10.1093/cz/zoy035/4983040 by guest on 13 July 2018 Ciach and Pe ˛ ksa Long-term changes in habitat use 7 Table 3. The results of multiple regression analysis testing for long-term changes in habitat use by the Tatra chamois R. rupicapra tatrica: correlation between year, population size, climatic conditions (temperature, precipitation, and snow depth), and within-a-year mean values of habitat variables recorded at the group sighting location (TNP; for parameters, see the “Materials and Methods” section; N ¼ 52) Dependent variable Effect Estimate SE 95% CL þ95% CL tP Effect size Altitude (mean) Intercept 4,707.94 852.26 2,992.44 6,423.45 5.52 0.000 Year 21.39 0.43 22.26 20.53 23.24 0.002 0.395 Population size 20.09 0.12 20.34 0.16 20.72 0.478 0.096 Temperature 13.64 4.39 4.79 22.48 3.10 0.003 0.404 Precipitation 0.05 0.19 20.34 0.43 0.25 0.803 0.126 Snow depth 0.06 0.37 20.68 0.79 0.16 0.873 0.161 Southerly aspect (mean) Intercept 274.92 81.84 110.18 439.65 3.36 0.002 Year 20.12 0.04 20.21 20.04 23.01 0.004 0.420 Population size 0.00 0.01 20.03 0.02 20.42 0.679 0.140 Temperature 20.25 0.42 21.10 0.60 20.59 0.558 0.159 Precipitation 20.02 0.02 20.06 0.02 21.17 0.248 0.061 Snow depth 0.04 0.04 20.03 0.11 1.25 0.219 0.169 Slope (mean) Intercept 46.25 28.90 211.92 104.42 1.60 0.116 Year 0.00 0.01 20.03 0.03 20.15 0.884 0.052 Population size 20.02 0.00 20.03 20.01 24.23 0.000 0.614 Temperature 0.22 0.15 20.08 0.52 1.49 0.142 0.106 Precipitation 0.00 0.01 20.01 0.01 0.11 0.912 0.075 Snow depth 0.00 0.01 20.02 0.03 0.40 0.693 0.138 Meadows (mean) Intercept 2114.53 110.03 2336.01 106.96 21.04 0.303 Year 0.07 0.06 20.04 0.19 1.33 0.191 0.200 Population size 0.03 0.02 0.00 0.07 2.17 0.035 0.387 Temperature 20.35 0.57 21.50 0.79 20.62 0.535 0.039 Precipitation 0.01 0.02 20.04 0.06 0.26 0.794 0.135 Snow depth 20.05 0.05 20.14 0.05 20.98 0.331 0.221 Trees and shrubs (mean) Intercept 236.77 60.72 2158.99 85.45 20.61 0.548 Year 0.02 0.03 20.04 0.08 0.66 0.514 0.048 Population size 0.01 0.01 20.01 0.02 0.63 0.533 0.118 Temperature 20.82 0.31 21.45 20.19 22.62 0.012 0.348 Precipitation 0.02 0.01 20.01 0.05 1.44 0.157 0.184 Snow depth 20.05 0.03 20.10 0.00 21.88 0.067 0.026 Distance to trail (mean) Intercept 4,557.58 1,104.35 2,334.65 6,780.52 4.13 0.000 Year 22.13 0.56 23.25 21.01 23.83 0.000 0.527 Population size 20.08 0.16 20.40 0.24 20.49 0.627 0.105 Temperature 4.28 5.69 27.18 15.74 0.75 0.456 0.078 Precipitation 20.13 0.25 20.63 0.37 20.52 0.604 0.046 Snow depth 0.21 0.47 20.74 1.16 0.44 0.659 0.025 Notes: Effect size is the z-transformed Pearson product-moment correlation coefﬁcient. Statistically signiﬁcant terms (P< 0.05) are shown in bold. or shrubs are also associated with group size: the larger the group, 2017). This allows them to utilize habitat with a potentially greater the more frequently it is found in such habitats. The antipredator risk of predation. Although group size generally increases with habi- benefits of group living, including dilution, satiation and confusion tat openness in large mammalian herbivores (Gerard and Loisel effects, vigilance, and selfish herding (Lehtonen and Jaatinen 2016) 1995), chamois co-occurring with lynx may adopt the opposite may explain such relationship. A larger group size improves the strategy. In open areas they likely have an advantage over their pri- chances of a potential threat being detected early, thereby reducing mary predators, since this type of habitats overlap with steep, rocky the risk of predation (Pe ´ rez-Barberı´a and Nores 1994). The preda- slopes, which are less suitable for successful hunting than habitats tors hunting chamois in the Tatras include the wolf Canis lupus, partially covered with dense forest vegetation. At present, habitat modification and fragmentation are among brown bear Ursus arctos, and lynx Lynx lynx (Jamrozy et al. 2007). The latter predator, which is known to be an important natural the main challenges facing animal populations (Fischer and threat to chamois (Molinari-Jobin et al. 2002), can use scrub for Lindenmayer 2007). With their unique ecological adaptations, high- concealment, from which an effective attack can be launched. In mountain species are particularly vulnerable to habitat change (Case et al. 2015). The effects of human activities in high-mountain such cases, the larger the number of herbivores in a group, the better they are able to monitor their surroundings and the earlier they can regions, be they local ones like sheep grazing and tourism, or global perceive a threat (Roberts 1996; Kluever et al. 2008; Beauchamp ones like climate change, can affect how animals use their habitats Downloaded from https://academic.oup.com/cz/advance-article-abstract/doi/10.1093/cz/zoy035/4983040 by guest on 13 July 2018 8 Current Zoology, 2018, Vol. 0, No. 0 Case MJ, Lawler JJ, Tomasevic JA, 2015. Relative sensitivity to climate and bring about long-term changes in their behavior. Although ani- change of species in northwestern North America. Biol Conserv 187: mals can adapt to functioning in dynamically changing habitats, 127–133. changes in species’ ecology are usually of a long-term nature and Chirichella R, Ciuti S, Apollonio M, 2013. Effects of livestock and non-native therefore hard to predict on the basis of short-term studies. This mouﬂon on use of high-elevation pastures by Alpine chamois. Mammal Biol work, the first documentation of long-term changes in habitat use 78:344–350. by ungulates, demonstrates that changes in the ecology of a species Chovancova ´ B, Zie ˛ ba F, Zwijacz-Kozica T, 2006. Polish and Slovakian count- can be induced by human activities: the abandonment of pasturage, ing of chamois: assumptions, methods and sources of errors. In: Krzan Z, high-mountain tourism, and climate change, over the time frame of editor. Tatrzanski Park Narodowy Na Tle Innych Go ´ rskich Tereno´w our study, have been the principal drivers of behavioral evolution in Chronionych, Tom II. Zakopane: TPN, 47–51. Chudı´k I, 1969. Ursachen der Verluste und der Einﬂuss der grossen Raubtiere the Tatra chamois. The present study also highlights the fact that auf die Population des Schalenwildes im Tatra-Nationalpark. 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Published: Apr 23, 2018
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