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Habitat selection of semi-free ranging European bison: Do bison preferred natural open habitats?

Habitat selection of semi-free ranging European bison: Do bison preferred natural open habitats? European bison (Bison bonasus) were successfully reintroduced in many free or semi-free areas across Europe during the last decades. Due to the increased numbers, the conflicts between human activities and bison are more frequent. Therefore the knowledge about spatial activity and habitat preference in new regions is the need for management decision making. We studied daily and seasonal habitat use of the semi-free European bison herd in the Czech Repub- lic from 2014 to 2019. The lead cows of the herd were collared with the GPS devices with a 30-minutes GPS fixes interval. The bison herd strongly preferred the managed open areas and supplementary feeding stations during the seasons (Jacob´s index from 0.49 to 0.99). On the contrary, they avoided the forest type and unmanaged open habitats (Jacob´s index from −0.23 to −0.69). The managed meadows and feeding places they used almost exclusively during the night while the forest during the day-light. Key words: habitat preference; Jacob´s index; forest, open areas; feeding station Editor: Bohdan Konôpka tion, the European bison should prevent the spread and 1. Introduction development of shrubs and trees and thus maintain these The European bison is the largest wild ungulate on the areas (Jirků & Dostál 2020). The presence of the Euro- European continent (Ramos et al. 2016). At the end of pean bison in these semi-wild farms has a significant the 19th century, it was on the verge of extinction, but effect on the species structure of habitats (Dostál et al. thanks to a successful reintroduction programme, more 2012). This is often the reason for the so-called rewild- than 40 wild yet geographically separated populations ing concept, which assumes the reintroduction of spe- currently live in Europe (Raczynski 2019). Due to the fact cies that have disappeared from the landscape or species that the European bison is introduced to new areas and farmed with the aim of increasing biodiversity without different types of environments, there is still a lack of a the need for active human management. This idea also unified view of its habitat preferences and what its natu- often includes the reintroduction of the European bison ral environment is. The European bison is considered a in many areas of Central and Eastern Europe, namely species that prefers a forest environment (Plumb et al. Germany, Poland, Lithuania and Latvia. Where several 2009; Bleier et al. 2012; Hofman-Kamińska & Kowalczyk highly suitable regions large enough to support a new 2012). However, the reason may be that the European herd were found, with the optimal areas being in Poland, bison is an example of a species that has been forced to the European bison was introduced in this way (Lord et move into the forest due to a combination of overgrowth al. 2020). Usually, semi-wild breeding is significantly of open areas after the last postglacial period and increas- limited (fenced) in space, however, with the ambition of ing human pressure on it (Cromssigt et al. 2012; Kerley spreading the European bison into the wild. And as like et al. 2012). Based on these assumptions and the fact in the case of open landscape, habitat preferences may that the European bison needs a large amount of her- be different. This can be reflected in the resulting impact baceous vegetation each day, in recent years the bison on different types of habitats. Therefore, we decided to has been very often introduced to places where it should serve as one of the species naturally farming open grassy evaluate the habitat preferences of European bison herds and bushy succession habitats. In these types of vegeta- bred in semi-wild breeding in the Czech Republic. *Corresponding author. Miloš Ježek, e-mail: jezekm@fld.czu.cz, phone: + 420 775 262 365 M. Zikmund et al. / Cent. Eur. For. J. 67 (2021) 30–34 The Jacob’s index was used to determine habitat pref- 2. Material and methods erences. The Jacob’s index was calculated according to 2.1. Study area the formula: D = (r – p)/(r + p – 2rp), Židlov is a nature reserve with an approximate area of 38 km and is one of the largest fields in Central Europe. It where r is the proportion of habitat used and p the proportion is located in the northern part of the Czech Republic and of habitat available. D varies from −1 (strong avoidance) to +1 was established in 2000 on the territory of a former mili- (strong preference), and values close to zero indicate that the tary training ground. 55% of the area is occupied by com- habitat is used in proportion to its availability (Jacobs 1974; mercial forests, of which 88% are conifers (pine, spruce) Kauhala and Aittilla 2008). and 12% deciduous trees (birch, oak). The rest of the area Data normality was tested using the one-sample (38%) consists of former impact areas that have been Kolmogorov-Smirnov test. If the distribution was not left to natural succession and currently form a forest- normal, a nonparametric test was used to compare the steppe landscape with a predominance of unmaintained data. For the Jacob’s index, 95% confidence intervals grasses, pioneer trees and shrubs. Cultivated agricultural were calculated to find the difference in different parts land occupies 5% and consists of maintained meadows of the year. used for grazing game. In the nature reserve there are The data were divided according to the time distri- red deer, fallow deer, mouflon, roe deer, and wild boar. bution in the year, either according to calendar months Since 2011, European bison, which have been imported or seasonality for spring (March, April, May), summer from Białowieża Forest and Kampinos National Park, (June, July, August), autumn (September, October, have been bred in the nature reserve. Currently, there November) and winter (December, January, February). is a herd of a total of 34 European bison in the nature The Oriana 4.02 software (Kovach Computing) and reserve (Raczyński 2019). Since 2014, wolves that are circular statistics were used to examine the distribution able to overcome fencing occasionally occur in the nature of habitat preference during the 24-hour cycle (Lehner reserve. 1996). Signic fi ant deviations from random distributions were investigated using the Rayleigh test of circular sta- tistics. 2.2. Data collection Between 2014 and 2018, the movement of the European 3. results bison herd was monitored with a telemetry collar (VEC- TRONIC Aerospace GmbH; type GPS Plus 5D). The On average across the entire time period, the European sampling frequency of GPS positions was 30 minutes bison herd used forest type (34%), feeding places (23%), (Červený et al. 2014). GPS data were used according to meadow (22%) and bush (21%). However, habitat use the method of Lewis et al. (2007), and all positions with varied significantly between seasons (Kruskal-Wallis DOP less than 6 were not used in the analyzes due to high Anova, p < 0.000; Fig. 1). Forest stands were mostly used inaccuracy of measurement. A total of 23,000 positions by bison in summer (47%) and least in autumn (20%), were available. feeding places mostly in winter (58%) and least in sum- Information about the environment comes from the mer (3%), meadows mostly in summer (33%) and least in mapping itself, when on the basis of field walks the area winter (5%), and bush most in summer (23%) and least was vectorized and habitats were divided into the follow- in winter (15%). ing main types: Meadow, Bush, Forest, Feeding places. 0 RGDH Z Meadow is a grassland economically maintained and %X VK R) HU VW restored, which is used to graze game kept in the nature ) LGHH JQ S DO FH reserve. Bush is formed by stands of early successional stages, which are gradually overgrown with shrubs and trees; these stands are not maintained. Forests are com- mercial forests used for wood production. Feeding places are places where supplementary feed (grain, corn, hay, haylage) is presented to game throughout the year. An area with a radius of 100 meters from the feeding facility was defined as a feeding place. :LQWHU 6SULQJ 6XPPHU $XWXPQ 2.3. Statistical analyses Occurrence and habitat data were processed in ArcGIS Fig. 1. Habitat utilization of bison during the seasons (% of to- tal location in the season) for different type of habitat (column 10.7 software (ESRI 2010). The spatial join tool was used = mean; whisker = mean ± 0.95 conf. interval). to link bison herd data and habitat type. 31 M. Zikmund et al. / Cent. Eur. For. J. 67 (2021) 30–34 As for the preferences in individual seasons (Fig. 2 index are greatest, European bison primarily use feed- and Table 1), the largest Jacob’s index showed a bison ing places and meadows during the night, while forest herd for feeding places in winter (0.98), spring (0.87), almost exclusively during the day. Bush, on the other autumn (0.86), only in summer the preference slightly hand, is used during dawn (Fig. 3). At the same time, decreased (to 0.41). Another type of vegetation that Euro- in meadows and forests there is a very high value of the pean bison strongly preferred were cultivated meadows. mean length of the vector and the concentration of data in In spring the Jacob’s index was 0.49, in summer 0.76, the preferred parts of the day throughout the year. With in autumn 0.70. Only in winter the index fell to −0.14. the exception of spring, the values of bushes are very low On the contrary, the European bison did not prefer, or and the significance is also an order of magnitude lower even deliberately avoided the forest and bush. The Jacob’s than for other types of stands. index for the forest ranged from −0.69 in the fall to −0.23 in the spring. It was similar for bush, when the index took values from −0.49 in winter to −0.26 in spring. &ŽƌĞƐ ƚ ƵƐŚ DĞĂĚŽǁ &ĞĞĚŝŶŐƉůĂĐ Ğ ʹϭ͘ Ϭ ʹϬ͘ ϱ Ϭ͘Ϭ Ϭ͘ϱ ϭ͘Ϭ ĞƚŶtŝ ƌ ƵƚƵŵ Ŷ ^ƵŵŵĞƌ Ŷŝƌ^Ɖ Ő Fig. 2. Habitat preference of bison during the seasons (Jacob´s index) for different type of habitat. Table 1. Values of Jacob´s index for month´s and different type of habitat. Fig. 3. Daily habitat selection during the spring: a) bush; b) Feeding place Meadow Bush Forest forest; c) feeding place; d) meadow. January 0.99 −0.42 −0.71 −0.67 February 0.96 −0.11 −0.23 −0.53 March 0.96 0.36 −0.36 −0.49 April 0.77 0.53 −0.28 −0.15 4. Discussion May 0.00 0.57 −0.27 −0.05 June 0.56 0.73 −0.22 −0.34 The high preference of the forest corresponds to the July -0.01 0.79 −0.31 −0.33 results found in other studies. As reported by Bleier et al. August 0.48 0.78 −0.24 −0.41 September 0.78 0.79 −0.21 −0.55 (2012) or Hofman-Kamińska and Kowalczyk (2012), the October 0.86 0.77 −0.17 −0.61 European bison prefer forest type environments, espe- November 0.94 0.72 −0.19 −0.73 December 0.99 0.02 −0.52 −0.75 cially during daytime (Červený et al. 2014; Marozas et al. 2019). Conversely, at night, European bison prefer open The distribution of habitat use also shows statisti- stands (Pucek et al. 2004; Daleszcyk et al. 2007; Marozas cally significant differences (Table 2). In winter, when et al. 2019). This corresponds to the theory that bison the differences in preference according to the Jacob’s changed their behaviour after the last postglacial period, Table 2. Circular statistics values for daily use of different type of habitat and season. Spring Summer Autumn Winter Mean vector (time) 01:00 21:33 12:23 00:08 Length of Mean Vector (r) 0.151 0.21 0.139 0.164 Feeding place Concentration 0.305 0.429 0.28 0.333 Rayleigh Test (p) < 0.001 < 0.001 < 0.001 < 0.001 Mean vector 00:35 0:58 00:22 02:33 Length of Mean Vector (r) 0.5423 0.486 0.412 0.200 Meadow Concentration 1.222 1.109 0.904 0.409 Rayleigh Test (p) < 0.001 < 0.001 < 0.001 < 0.001 Mean vector 12:27 12:42 12:06 12:04 Length of Mean Vector (r) 0.338 0.459 0.509 0.464 Forest Concentration 0.719 1.033 1.179 1.046 Rayleigh Test (p) < 0.001 < 0.001 < 0.001 < 0.001 Mean vector 00:11 00:06 20:37 19:14 Length of Mean Vector (r) 0.299 0.099 0.035 0.092 Bush Concentration 0.628 0.199 0.069 0.184 Rayleigh Test (p) < 0.001 < 0.01 < 0.01 < 0.01 32 M. Zikmund et al. / Cent. Eur. For. J. 67 (2021) 30–34 which was associated with a decrease in open areas and 5. Conclusion increasing activity of people for whom European bison We conclude that bison highly preferred the managed were prey (Cromssigt et al. 2012; Kerley et al. 2012). open habitats (meadows) and feeding sites all year Their frequent stay in open areas during the night can around. The forest habitats bison use usually during the also be the result of better grazing, when, like other rumi- daylight which is related to antipredation behavior. In nants, they look for richer food habitats at a time when contrary they avoid the opened unmanaged succession they are less disturbed by humans (Whittaker & Knight areas. This can cause conflicts between the humans and 1998). This is also confirmed by conclusions about the bison interests. It´s also necessary to consider the rela- type of habitat preference. In most cases within our study tionship with other game species because of the high these were maintained grasslands, regularly managed. preference for feeding stations by bison. This can cause Their share in the total area is only 5%, but bison occur that other species will have limited access to these places. in it up to 33% of the total time of day (Summer). Other In the case of bison reintroduction in areas with natural successive shrub formations (Bush) are avoided intensively cultivated agricultural areas, it is appropri- throughout the year, even though they make up more ate to cultivate a special pasture areas for it and thus than 43% of the total area. On average, they spend only eliminate possible migration outside the core area and 21% of their total time on them. In other studies, Euro- prevent damages and conflicts. A very common goal of pean bison similarly preferred cultivated farmland. E.g. bison reintroduction is to eliminate woody plants and in Lithuania (Marozas et al. 2019), European bison spent other aggressive species of plant on the open succession up to 56% of their time at night there, of which only 7% areas by bison grazing. However, this purpose proves on the uncultivated land to which we can compare bush. to be unsuitable for large semi free areas or in the free- On the contrary, they signic fi antly preferred qualitatively range areas. It is suitable to implement it only in small rich types of vegetation, such as rape or cereals. European limited farms, where the bison has a limited choice of bison living on the borders of the Białowieża National habitat and food sources. To establish the management Park showed similar preferences, when their home dis- plans and true decision making policy, a detailed study of tricts intervened in the agricultural landscape. This also habitat preferences is suitable, especially in areas newly supports the n fi dings of Mendoza and Palmquist (2008), inhabited by bison. who rank the bison morphologically among the species adapted to live in rich grass communities. On the con- trary, the results of food analyses carried out in the cen- Acknowledgements tral part of the Białowieża National Park, where neither feeding is carried out nor cultivated agricultural crops are This study was supported by grants QK1910462 of the Ministry present, suggest that European bison prefer highly nutri- of Agriculture and B_19_02 of FFWS. tious and easily digestible parts of woody plants, shrubs and herbs to grasses (Kowalczyk et al. 2019). At the same time, they also point out that the content of graminoids references (which contain a large percentage of grasses) was higher, Bleier, N., Lehoczki, N., Újváry, D., Szemethy, L., Csány, especially in the period when the supply of quality trees S., 2012: Relationships between wild ungulates den- and shrubs was declining. This indicates a high degree sity and crop damage in Hungary. Acta Theriologica, of plasticity of the bison in food selection and they can 57:351–359. thus be described as a generalist (Freeland 1991), where Cromsigt, J. P. G. M., Kerley, G. I. H., Kowalczyk, R., this type of animal prefers high quality food and is only 2012: The difficulty of using species distribution selective if its presence in the environment is high. modeling for the conservation of refugee species–the The theory of the European bison as a generalist is example of European bison. Diversity and Distribu- also supported by the high intensity of the use of feeding tions,18:1253–1257. places, which in our study achieved very high preferences Červený, J., Ježek, M., Holá, M., Zikmund, M., Kušta, T., (Winter Jacob’s index = 1), and where the bison spent up Hanzal, V., Kropil, R., 2014: Daily activity rhythm to 60% of the day in their immediate vicinity (< 100 m). and habitat use of the semi-free European bison herd Such a high preference for feeding sites can cause a during the growing season. Lesnícky časopis – For- significant reduction in flight distance and also to attrac- estry Journal, 60:199–204. tion by strengthening an animal’s behavior through posi- Daleszczyk, K., Krasińska, M., Krasiński, Z. A., Bune- tive reinforcement and encouraging movement towards a vich, A. N., 2007: Habitat structure, climatic factors, stimulus (Haidt et al. 2018). Bisons aggregation by feed- and habitat use by European bison (Bison bonasus) ing sites may facilitate transmission of nematode Ash- in Polish and Belarussian parts of the Białowiez˙ a worthius sidemi, which occurs by ingesting an invasive Forest, Poland. Canadian Journal of Zoology, 85: larval stage (Radwan et al. 2010; Vadlejch et al. 2017). 261–272. 33 M. Zikmund et al. / Cent. Eur. For. J. 67 (2021) 30–34 Dostál, D., Jirků, M., Konvička, M., Čížek, L., Šálek, M., Lord, C. M., Wirebach, K. P., Tompkins, J., Bradshaw- 2012: Návrat zubra evropského (Bison bonasus) do Wilson, C., Shaffer, Ch. L., 2020: Reintroduction of České republiky. Česká krajina, Kutná Hora, 120 p. the European bison (Bison bonasus) in central-east- Freeland, W. J., 1991: Plant secondary metabolites: bio- ern Europe: a case study. International Journal of chemical coevolution with herbivores. Plant defen- Geographical Information Science, 34:1628–1647. ses against mammalian herbivory. CRC Press, Boca Marozas, V., Kibiša, A., Brazaitis, G., Jõgiste, K., Raton, p. 61–81. Šimkevičius, K., Bartkevičius, E., 2019: Distribu- Haidt, A., Kamiński, T., Borowik, T., Kowalczyk, R., tion and Habitat Selection of Free-Ranging European 2018: Human and the beast – Flight and aggressive Bison (Bison bonasus L.) in a Mosaic Landscape–A responses of European bison to human disturbance. Lithuanian Case. Forests, 10:339–345. PLOS ONE, 13:e0200635. Mendoza, M., Palmqvist, P., 2008: Hypsodonty in ungu- Hofman-Kamińska, E., Kowalczyk, R., 2012: Farm lates: an adaptation for grass consumption or for Crops Depre-dation by European Bison (Bison bona- foraging in open habitat? Journal of Zoology, 274: sus) in the Vicinity of Forest Habitats in Northeastern 134–142. Poland. Environmental Management, 50:530–541. Plumb, G. E., White, P. J., Coughenour, M. B., Wallen, R. Jacobs, J., 1974:. Quantitative measurement of food sele- L., 2009: Carrying capacity, migration, and dispersal ction. Oecologia, 14:413–417. in Yellowstone bison. Biologicall Conservation, 142: Jirků, M., Dostál, D., 2020: Zhodnocení managementu 2377–2387. bezlesí VVP Milovice – Mladá přirozenou pastvou vel- Pucek, Z., Belousova, I. P., Krasińska, M., Krasiński, Z. kých kopytníků 2015 – 2019. Praha, 25 p. A., Olech, W., 2004: European bison. Status survey Kauhala, K., Auttila, M., 2010: Estimating habi - and conservation action plan. IUCN/SSC Bison Spe- tat selection of badgers-a test between different cialist Group. IUCN, Gland, Switzerland. methods. Journal of Vertebrate Biology, 59:16–25. Raczyński, J., 2019: European bison pedigree book. Kerley, G. I. H., Kowalczyk, R., Cromsigt, J. P. G. M. Białowieski Park Narodowy, 100 p. 2012: Conservation implications of the refugee spe- Radwan, J., Demiaszkiewicz, A. W., Kowalczyk, R., cies concept and the European bison: king of the for- Lachowicz, J., Kawalko, A., Wojcik, J. M.et al., est or refugee in a marginal habitat? Ecography, 35: 2010: An evaluation of two potential risk factors, 519–529 MHC diversity and host density, for infection by an Kowalczyk, R., Wójcik, J. M., Taberlet, P., Kamiński, T., invasive nematode Ashworthius sidemi in endangered Miquel, Ch., Valentini, A. et al., 2019: Foraging plas- European bison (Bison bonasus). Biological Conser- ticity allows a large herbivore to persist in a sheltering vation, 143:2049–2053. forest habitat: DNA metabarcoding diet analysis of Ramos, A., Petit, O., Longour, P., Pascuarett, C., Sueur, the European bison. Forest Ecology and Manage- S., 2015: Collective decision making during group ment, 449:117474. movements in Europea bison (Bison bonasus). Ani- Lehner, P. N., 1996: Spatial orientation and time: circu- mal Behavior, 109:149–160. lar statistics and spatial patterns. In: Lehner, P. N. Ramos, A., Petit, O., Longour, P., Pasquaretta, C., (ed.): Handbook of ethological methods. Cambridge Sueur, C., 2016: Space use and movement patterns University Press, p. 485–521. in a semi-free-ranging herd of European bison (Bison Lewis, J. S., Rachlow, J. L., Garton, E. O., Vierling, L. A., bonasus). PloS One, 11:e0147404. 2007: Effects of habitat on GPS collar performance: Vadlejch, J., Kyriánová, I. A., Rylková, K., Zikmund, M., using data screening to reduce location error. Journal Langrová, I., 2017: Did European bison conservation of Apply Ecology, 44:663–671. program introduced an alien parasite to the Czech Republic? Biological Invasions, 19:1121–1125. Whittaker, D., Knight, R. L., 1998: Understanding wild- life responses to humans. Wildlife Society Bulletin, 26:312–317. http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Forestry Journal de Gruyter

Habitat selection of semi-free ranging European bison: Do bison preferred natural open habitats?

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Abstract

European bison (Bison bonasus) were successfully reintroduced in many free or semi-free areas across Europe during the last decades. Due to the increased numbers, the conflicts between human activities and bison are more frequent. Therefore the knowledge about spatial activity and habitat preference in new regions is the need for management decision making. We studied daily and seasonal habitat use of the semi-free European bison herd in the Czech Repub- lic from 2014 to 2019. The lead cows of the herd were collared with the GPS devices with a 30-minutes GPS fixes interval. The bison herd strongly preferred the managed open areas and supplementary feeding stations during the seasons (Jacob´s index from 0.49 to 0.99). On the contrary, they avoided the forest type and unmanaged open habitats (Jacob´s index from −0.23 to −0.69). The managed meadows and feeding places they used almost exclusively during the night while the forest during the day-light. Key words: habitat preference; Jacob´s index; forest, open areas; feeding station Editor: Bohdan Konôpka tion, the European bison should prevent the spread and 1. Introduction development of shrubs and trees and thus maintain these The European bison is the largest wild ungulate on the areas (Jirků & Dostál 2020). The presence of the Euro- European continent (Ramos et al. 2016). At the end of pean bison in these semi-wild farms has a significant the 19th century, it was on the verge of extinction, but effect on the species structure of habitats (Dostál et al. thanks to a successful reintroduction programme, more 2012). This is often the reason for the so-called rewild- than 40 wild yet geographically separated populations ing concept, which assumes the reintroduction of spe- currently live in Europe (Raczynski 2019). Due to the fact cies that have disappeared from the landscape or species that the European bison is introduced to new areas and farmed with the aim of increasing biodiversity without different types of environments, there is still a lack of a the need for active human management. This idea also unified view of its habitat preferences and what its natu- often includes the reintroduction of the European bison ral environment is. The European bison is considered a in many areas of Central and Eastern Europe, namely species that prefers a forest environment (Plumb et al. Germany, Poland, Lithuania and Latvia. Where several 2009; Bleier et al. 2012; Hofman-Kamińska & Kowalczyk highly suitable regions large enough to support a new 2012). However, the reason may be that the European herd were found, with the optimal areas being in Poland, bison is an example of a species that has been forced to the European bison was introduced in this way (Lord et move into the forest due to a combination of overgrowth al. 2020). Usually, semi-wild breeding is significantly of open areas after the last postglacial period and increas- limited (fenced) in space, however, with the ambition of ing human pressure on it (Cromssigt et al. 2012; Kerley spreading the European bison into the wild. And as like et al. 2012). Based on these assumptions and the fact in the case of open landscape, habitat preferences may that the European bison needs a large amount of her- be different. This can be reflected in the resulting impact baceous vegetation each day, in recent years the bison on different types of habitats. Therefore, we decided to has been very often introduced to places where it should serve as one of the species naturally farming open grassy evaluate the habitat preferences of European bison herds and bushy succession habitats. In these types of vegeta- bred in semi-wild breeding in the Czech Republic. *Corresponding author. Miloš Ježek, e-mail: jezekm@fld.czu.cz, phone: + 420 775 262 365 M. Zikmund et al. / Cent. Eur. For. J. 67 (2021) 30–34 The Jacob’s index was used to determine habitat pref- 2. Material and methods erences. The Jacob’s index was calculated according to 2.1. Study area the formula: D = (r – p)/(r + p – 2rp), Židlov is a nature reserve with an approximate area of 38 km and is one of the largest fields in Central Europe. It where r is the proportion of habitat used and p the proportion is located in the northern part of the Czech Republic and of habitat available. D varies from −1 (strong avoidance) to +1 was established in 2000 on the territory of a former mili- (strong preference), and values close to zero indicate that the tary training ground. 55% of the area is occupied by com- habitat is used in proportion to its availability (Jacobs 1974; mercial forests, of which 88% are conifers (pine, spruce) Kauhala and Aittilla 2008). and 12% deciduous trees (birch, oak). The rest of the area Data normality was tested using the one-sample (38%) consists of former impact areas that have been Kolmogorov-Smirnov test. If the distribution was not left to natural succession and currently form a forest- normal, a nonparametric test was used to compare the steppe landscape with a predominance of unmaintained data. For the Jacob’s index, 95% confidence intervals grasses, pioneer trees and shrubs. Cultivated agricultural were calculated to find the difference in different parts land occupies 5% and consists of maintained meadows of the year. used for grazing game. In the nature reserve there are The data were divided according to the time distri- red deer, fallow deer, mouflon, roe deer, and wild boar. bution in the year, either according to calendar months Since 2011, European bison, which have been imported or seasonality for spring (March, April, May), summer from Białowieża Forest and Kampinos National Park, (June, July, August), autumn (September, October, have been bred in the nature reserve. Currently, there November) and winter (December, January, February). is a herd of a total of 34 European bison in the nature The Oriana 4.02 software (Kovach Computing) and reserve (Raczyński 2019). Since 2014, wolves that are circular statistics were used to examine the distribution able to overcome fencing occasionally occur in the nature of habitat preference during the 24-hour cycle (Lehner reserve. 1996). Signic fi ant deviations from random distributions were investigated using the Rayleigh test of circular sta- tistics. 2.2. Data collection Between 2014 and 2018, the movement of the European 3. results bison herd was monitored with a telemetry collar (VEC- TRONIC Aerospace GmbH; type GPS Plus 5D). The On average across the entire time period, the European sampling frequency of GPS positions was 30 minutes bison herd used forest type (34%), feeding places (23%), (Červený et al. 2014). GPS data were used according to meadow (22%) and bush (21%). However, habitat use the method of Lewis et al. (2007), and all positions with varied significantly between seasons (Kruskal-Wallis DOP less than 6 were not used in the analyzes due to high Anova, p < 0.000; Fig. 1). Forest stands were mostly used inaccuracy of measurement. A total of 23,000 positions by bison in summer (47%) and least in autumn (20%), were available. feeding places mostly in winter (58%) and least in sum- Information about the environment comes from the mer (3%), meadows mostly in summer (33%) and least in mapping itself, when on the basis of field walks the area winter (5%), and bush most in summer (23%) and least was vectorized and habitats were divided into the follow- in winter (15%). ing main types: Meadow, Bush, Forest, Feeding places. 0 RGDH Z Meadow is a grassland economically maintained and %X VK R) HU VW restored, which is used to graze game kept in the nature ) LGHH JQ S DO FH reserve. Bush is formed by stands of early successional stages, which are gradually overgrown with shrubs and trees; these stands are not maintained. Forests are com- mercial forests used for wood production. Feeding places are places where supplementary feed (grain, corn, hay, haylage) is presented to game throughout the year. An area with a radius of 100 meters from the feeding facility was defined as a feeding place. :LQWHU 6SULQJ 6XPPHU $XWXPQ 2.3. Statistical analyses Occurrence and habitat data were processed in ArcGIS Fig. 1. Habitat utilization of bison during the seasons (% of to- tal location in the season) for different type of habitat (column 10.7 software (ESRI 2010). The spatial join tool was used = mean; whisker = mean ± 0.95 conf. interval). to link bison herd data and habitat type. 31 M. Zikmund et al. / Cent. Eur. For. J. 67 (2021) 30–34 As for the preferences in individual seasons (Fig. 2 index are greatest, European bison primarily use feed- and Table 1), the largest Jacob’s index showed a bison ing places and meadows during the night, while forest herd for feeding places in winter (0.98), spring (0.87), almost exclusively during the day. Bush, on the other autumn (0.86), only in summer the preference slightly hand, is used during dawn (Fig. 3). At the same time, decreased (to 0.41). Another type of vegetation that Euro- in meadows and forests there is a very high value of the pean bison strongly preferred were cultivated meadows. mean length of the vector and the concentration of data in In spring the Jacob’s index was 0.49, in summer 0.76, the preferred parts of the day throughout the year. With in autumn 0.70. Only in winter the index fell to −0.14. the exception of spring, the values of bushes are very low On the contrary, the European bison did not prefer, or and the significance is also an order of magnitude lower even deliberately avoided the forest and bush. The Jacob’s than for other types of stands. index for the forest ranged from −0.69 in the fall to −0.23 in the spring. It was similar for bush, when the index took values from −0.49 in winter to −0.26 in spring. &ŽƌĞƐ ƚ ƵƐŚ DĞĂĚŽǁ &ĞĞĚŝŶŐƉůĂĐ Ğ ʹϭ͘ Ϭ ʹϬ͘ ϱ Ϭ͘Ϭ Ϭ͘ϱ ϭ͘Ϭ ĞƚŶtŝ ƌ ƵƚƵŵ Ŷ ^ƵŵŵĞƌ Ŷŝƌ^Ɖ Ő Fig. 2. Habitat preference of bison during the seasons (Jacob´s index) for different type of habitat. Table 1. Values of Jacob´s index for month´s and different type of habitat. Fig. 3. Daily habitat selection during the spring: a) bush; b) Feeding place Meadow Bush Forest forest; c) feeding place; d) meadow. January 0.99 −0.42 −0.71 −0.67 February 0.96 −0.11 −0.23 −0.53 March 0.96 0.36 −0.36 −0.49 April 0.77 0.53 −0.28 −0.15 4. Discussion May 0.00 0.57 −0.27 −0.05 June 0.56 0.73 −0.22 −0.34 The high preference of the forest corresponds to the July -0.01 0.79 −0.31 −0.33 results found in other studies. As reported by Bleier et al. August 0.48 0.78 −0.24 −0.41 September 0.78 0.79 −0.21 −0.55 (2012) or Hofman-Kamińska and Kowalczyk (2012), the October 0.86 0.77 −0.17 −0.61 European bison prefer forest type environments, espe- November 0.94 0.72 −0.19 −0.73 December 0.99 0.02 −0.52 −0.75 cially during daytime (Červený et al. 2014; Marozas et al. 2019). Conversely, at night, European bison prefer open The distribution of habitat use also shows statisti- stands (Pucek et al. 2004; Daleszcyk et al. 2007; Marozas cally significant differences (Table 2). In winter, when et al. 2019). This corresponds to the theory that bison the differences in preference according to the Jacob’s changed their behaviour after the last postglacial period, Table 2. Circular statistics values for daily use of different type of habitat and season. Spring Summer Autumn Winter Mean vector (time) 01:00 21:33 12:23 00:08 Length of Mean Vector (r) 0.151 0.21 0.139 0.164 Feeding place Concentration 0.305 0.429 0.28 0.333 Rayleigh Test (p) < 0.001 < 0.001 < 0.001 < 0.001 Mean vector 00:35 0:58 00:22 02:33 Length of Mean Vector (r) 0.5423 0.486 0.412 0.200 Meadow Concentration 1.222 1.109 0.904 0.409 Rayleigh Test (p) < 0.001 < 0.001 < 0.001 < 0.001 Mean vector 12:27 12:42 12:06 12:04 Length of Mean Vector (r) 0.338 0.459 0.509 0.464 Forest Concentration 0.719 1.033 1.179 1.046 Rayleigh Test (p) < 0.001 < 0.001 < 0.001 < 0.001 Mean vector 00:11 00:06 20:37 19:14 Length of Mean Vector (r) 0.299 0.099 0.035 0.092 Bush Concentration 0.628 0.199 0.069 0.184 Rayleigh Test (p) < 0.001 < 0.01 < 0.01 < 0.01 32 M. Zikmund et al. / Cent. Eur. For. J. 67 (2021) 30–34 which was associated with a decrease in open areas and 5. Conclusion increasing activity of people for whom European bison We conclude that bison highly preferred the managed were prey (Cromssigt et al. 2012; Kerley et al. 2012). open habitats (meadows) and feeding sites all year Their frequent stay in open areas during the night can around. The forest habitats bison use usually during the also be the result of better grazing, when, like other rumi- daylight which is related to antipredation behavior. In nants, they look for richer food habitats at a time when contrary they avoid the opened unmanaged succession they are less disturbed by humans (Whittaker & Knight areas. This can cause conflicts between the humans and 1998). This is also confirmed by conclusions about the bison interests. It´s also necessary to consider the rela- type of habitat preference. In most cases within our study tionship with other game species because of the high these were maintained grasslands, regularly managed. preference for feeding stations by bison. This can cause Their share in the total area is only 5%, but bison occur that other species will have limited access to these places. in it up to 33% of the total time of day (Summer). Other In the case of bison reintroduction in areas with natural successive shrub formations (Bush) are avoided intensively cultivated agricultural areas, it is appropri- throughout the year, even though they make up more ate to cultivate a special pasture areas for it and thus than 43% of the total area. On average, they spend only eliminate possible migration outside the core area and 21% of their total time on them. In other studies, Euro- prevent damages and conflicts. A very common goal of pean bison similarly preferred cultivated farmland. E.g. bison reintroduction is to eliminate woody plants and in Lithuania (Marozas et al. 2019), European bison spent other aggressive species of plant on the open succession up to 56% of their time at night there, of which only 7% areas by bison grazing. However, this purpose proves on the uncultivated land to which we can compare bush. to be unsuitable for large semi free areas or in the free- On the contrary, they signic fi antly preferred qualitatively range areas. It is suitable to implement it only in small rich types of vegetation, such as rape or cereals. European limited farms, where the bison has a limited choice of bison living on the borders of the Białowieża National habitat and food sources. To establish the management Park showed similar preferences, when their home dis- plans and true decision making policy, a detailed study of tricts intervened in the agricultural landscape. This also habitat preferences is suitable, especially in areas newly supports the n fi dings of Mendoza and Palmquist (2008), inhabited by bison. who rank the bison morphologically among the species adapted to live in rich grass communities. On the con- trary, the results of food analyses carried out in the cen- Acknowledgements tral part of the Białowieża National Park, where neither feeding is carried out nor cultivated agricultural crops are This study was supported by grants QK1910462 of the Ministry present, suggest that European bison prefer highly nutri- of Agriculture and B_19_02 of FFWS. tious and easily digestible parts of woody plants, shrubs and herbs to grasses (Kowalczyk et al. 2019). At the same time, they also point out that the content of graminoids references (which contain a large percentage of grasses) was higher, Bleier, N., Lehoczki, N., Újváry, D., Szemethy, L., Csány, especially in the period when the supply of quality trees S., 2012: Relationships between wild ungulates den- and shrubs was declining. This indicates a high degree sity and crop damage in Hungary. 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Journal

Forestry Journal – de Gruyter

Published: Mar 1, 2021

Keywords: habitat preference; Jacob´s index; forest; open areas; feeding station

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