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Article

European Beech Masting Cycles and the Spatial Distribution of Wisents in the Bieszczady Mountains, Poland

by
Aleksandra Wołoszyn-Gałęza
1,
Maciej Januszczak
1 and
Kajetan Perzanowski
2,*
1
Carpathian Wildlife Research Station, Museum and Institute of Zoology Polish Academy of Sciences, 38-700 Ustrzyki Dolne, Poland
2
Institute of Biological Sciences, Catholic University of Lublin, 20-708 Lublin, Poland
*
Author to whom correspondence should be addressed.
Forests 2024, 15(9), 1618; https://doi.org/10.3390/f15091618
Submission received: 10 July 2024 / Revised: 10 September 2024 / Accepted: 13 September 2024 / Published: 13 September 2024
(This article belongs to the Section Forest Ecology and Management)
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Abstract

:
The variability of food resources considerably affects the habitat preferences of animals. In mast years, the availability of highly nutritive food increases significantly. We tested whether changes in the distribution of the areas of wisents, Bison bonasus L. concentration, in the Bieszczady Mountains, Poland, were connected with the availability of beechnuts. In the two beech masting years of 2013 and 2022, we considered the months with the highest availability of beechnuts, namely September and October. The beechnut crop varied significantly between as little as 2.4 g dry matter/m2, recorded within the Baligród herd’s home range in 2013, and up to 238.8 g dry matter/m2 within the Tworylne herd’s range, also in 2013. The analysis of the spatial distribution of beech stands within various parts of the wisents’ home range showed that within the 95% kernel area, their share was mostly high, varying between 25.7% and 42.8%. Meanwhile, within the 50% kernel area, it was generally much lower, except for the year 2022 in the case of the Baligród herd. The densities of wisents varied significantly between the kernel areas of 95% and 50%, ranging between 0.04 and 0.08 animals/ha and 0.17 and 0.48 animals/ha, respectively. However, there was no statistical difference between the figures for all plots tested within the home range of the wisent population and plots dominated by beech. Based on the data obtained in this study, habitat selection patterns of wisents in the Bieszczady Mountains cannot be explained by the availability of beech stands and the phenomenon of mast years.

1. Introduction

Temporally variable seed production, known as masting, is typical for many tree species, e.g., European beech, Fagus sylvatica L. (Fagales: Fagaceae) or the pedunculate oak, Quercus robur L. (Fagales: Fagaceae). This phenomenon may have strong effect upon foraging conditions for many forest-dwelling species including rodents, ungulates or birds [1,2,3].
In the case of the oak, a massive crop of acorns occurs every 2–3 years, but the European beech usually produces seeds in 5–8-year and sometimes even in 10–11-year intervals [4,5,6,7,8]. Such temporal patterns dramatically affect the availability of highly nutritive forage for forest animals, which in consecutive years may change from several kilograms per sq. meter to zero [9,10]. Therefore, animal population dynamics, spatial distribution, migrations and even survival frequently correlate with mast years [1,11,12,13,14].
It has been found that fluctuations in seed production in European beech are connected with cold and wet summers occurring two years before the masting event, followed by hot and dry summers the next year [15]. However, a high amount of rainfall in spring during the beech flowering period, may negatively influence the amount of beechnuts produced, probably due to its interference with the pollination process [16,17]. Hence, long-term fluctuation patterns of weather conditions have a prevailing effect upon beechnut crops [2,18]. Among the other factors that may influence the masting patterns are the age of tree stand, its canopy closure and the elevation above sea level of a given site [19,20,21].
Large quantities of tree seeds that suddenly appear, like acorns or beechnuts, are highly attractive not only to small rodents or birds but also to a number of large mammals like wild boars, bears or various ungulate species [2,14,22,23,24]. Therefore, we could expect that during September–October in mast years, wisents will tend to concentrate in tree stands dominated by beech trees.
In this study, we compared changes in the distribution of the areas of European bison or wisents, Bison bonasus L. (Artiodactyla: Bovidae) concentration with the spatial distribution of the beechnut crops, in the Bieszczady Mountains, during the months with the highest availability of beechnuts (September–October). The years of beech masting studied were 2013 and 2022, in addition to November 2012–August 2013, and November 2021–August 2022 (periods when beechnuts were not available). Our aim was to test the hypothesis and determine whether the availability of beechnuts may temporally significantly influence the habitat use of wisents.

2. Study Area, Material and Methods

We estimated the beechnut crop in three forest districts of the Bieszczady Mountains (southeastern Poland): Baligród, Komańcza and Lutowiska. Beechnut sampling was performed in 2013 and in 2022, i.e., in only two mast years occurring during the studied period. In 2013, 5 sampling plots were established in Komańcza, 19 plots in Baligród and 6 in Lutowiska. In 2022, there were 6 plots in Komańcza, 16 in Baligród and 8 in Lutowiska. The plots were established randomly within mature beech stands. Due to logging activities, in some cases, it was not possible to sample the same plots in both years. Therefore, following GPS readings, we sampled sites as close to the original locations as possible. The forest districts of Baligród and Komańcza are inhabited by the western subpopulation of B. bonasus in the Bieszczady Mountains (henceforth called the Baligród herd). The Lutowiska District is within the home range of the eastern subpopulation (henceforth called the Tworylne herd) [25] (Figure 1).
Plots were established in stands dominated by F. sylvatica aged between 57 and 124 years (according to standard forestry maps), in most cases, at the same sites or possibly the nearest location, in 2013 and 2022, according to the GPS Garmin E-trex Vista (Garmin International Corp., Olathe, KS, USA). The dimensions of randomly allocated quadrate plots were 1 × 1 m. Within the plot areas, all beechnuts were collected from the ground. Samples were weighted with an accuracy of 0.1 g to obtain the value of biomass and subsequently dried four times at 70 °C for 12 h to obtain their dry mass.
The elevation above sea level for the tested sites was determined with the GPS Garmin E-trex Vista (Garmin International Corp., Olathe, KS, USA).
Within the home ranges of both monitored wisent herds, beech stands were, on average, situated at higher elevations compared to other types of tree stands. Those differences ranged from 27 to 66 m and 89 to 94 m within the 50% and 95% kernel areas for the Baligród herd, respectively, and from 86 to 95 m and 70 to 75 m within the 50% and 95% kernel areas for the Tworylne herd, respectively. The maximal elevations recorded for beech stands were 746 m (SD= 132) above sea level within the 95% kernel area of the Baligród herd and 665 m (SD = 103) for 95% kernel area of the Tworylne herd.
Presence records of wisents were obtained during the routine monitoring program of the wisent population inhabiting the Bieszczady Mountains, performed by the staff of the Carpathian Wildlife Research Station of the Polish Academy of Sciences (CWRS PAS) in cooperation with the Forest Service. All signs of the presence of wisents (visual observations, tracts, feces) as well as data obtained through the radiotracking of several individuals fitted with radiocollars were considered. In both years, the spatial distributions of both wisent herds were determined for the 2 months with the highest availability of beechnuts (September and October) and separately for the remaining part of the year (November–August) [26].
The population numbers of wisents in the Bieszczady Mountains changed considerably between 2013 and 2022 from 270 to 729 individuals (database of CWRS PAS).
The analysis of the spatial distribution of the wisent population was carried out with ArcView 9.3 (Esri Polska: https://www.esri.pl/) (accessed on 18 October 2022).
The calculation of the home range area (95% kernel) and concentration sites (50% kernel) was performed according to Worton [27].
The differences among the amounts of beechnut crop, wisent densities and the proportion of beech stands were estimated with Z test, using software PQStat v.1.8.6 (PQStat software: https://pqstat.pl/) (accessed on 18 October 2022).

3. Results

The lowest value of the beechnut crop (2.4 g dry matter/m2) was recorded within the Baligród herd home range in 2013. The highest value of the beechnut crop (238.8 g dry matter/m2) was found within the Tworylne herd range, also in 2013. Generally, however, the higher values of the beechnut crop in both sites were found in 2013. In 2013 and 2022, the beechnut crop was higher within the home range of the Tworylne herd. Nevertheless, a comparison of those values did not show statistically significant differences (0.4032, p ≤ 0.05) (Table 1).
The analysis of the spatial distribution of beech stands within various parts of the home range of wisents shows that within the 95% kernel area, their proportion was rather high, varying between 26% and 43%. Meanwhile, within the 50% kernel area, it was generally much lower (mostly below 10%), except in 2022 in the case of the Baligród herd. Also, during the remaining part of the year (November–August), the percentage of beech stands within the whole population range was significantly higher that within the concentrated areas of wisents (Table 2).
The densities of wisents varied significantly between the 95% and 50% kernel areas, ranging between 0.04 and 0.08 animals/ha and 0.17 and 0.48 animals/ha, respectively. However, there was no statistical difference in the recorded wisent presence between the figures for all plots within the home range of the population and plots dominated by beech (Table 3).
Those data were compared with the spatial distribution of the wisent herds during the remaining months of the same years (Table 4).

4. Discussion

The phenomenon of masting is known to play a significant role in the life cycles of many animal species like rodents, small birds, wild boars or black bears [1,2,10,12,14,23]. Wisents inhabiting the Białowieska Forest are known to intensively feed on acorns [28]. Since the home ranges of the wisent population in the Bieszczady Mountains are covered by a large proportion of beech stands, beech masting may influence the temporal patterns of spatial distribution of this population through the temporal availability of high-quality forage.
Occasional observations carried out by foresters in the mast years in the BieszczadyMountains report a frequent presence of wisents during the autumn in beech stands. The biomass of the ground flora and browse available within those habitats in this season is marginal [29,30,31]. Therefore, a potentially attracting factor for wisents could be the sudden accessibility of high-energy beechnuts.
However, our data show that the proportion of tree stands dominated by beech within the 50% kernel areas (i.e., the concentration sites of wisents) was generally low, much lower than within the whole 95 % kernel area, and this difference was significant. This ratio was similar in the months with high availability of beechnuts and in the remaining part of the year (Table 2). Therefore, the presence of beech trees could not explain the concentration of wisents within the 50% kernel areas.
Comparison of the densities of wisents during the mast years and seasons with no or low accessibility of beechnuts (Table 3 and Table 4) show that some other factors may play a main role in the selection of concentration sites by those animals. In some sites, wisents could be disturbed by the hunting season for red deer. Also, weather conditions could be unfavorable, especially at higher elevations that were dominated by pure beech stands, like early snow fall at that time [32,33,34]. Nevertheless, we had no access to such data. Hence, it is difficult to explain such observations, like the case of the Tworylne herd, where densities of wisents in the forest stands dominated by beech during the years with low accessibility of beechnuts highly exceeded the figures from the mast years.
Therefore, although wisents undoubtedly willingly consume available beechnuts, their habitat selection patterns could not be explained on the basis of available data by the phenomenon of mast years. This could be a result of a generally high availability of beech in this region, very abundant ground flora [31] and a coexistence of other factors influencing the spatial distribution of those animals, which should be a topic for future studies.

5. Conclusions

Since the vegetative period of plants ends in autumn, the natural food supply for ruminants becomes substantially limited in this season. Therefore, cyclically occurring beechnut crops may potentially become an additional, high-energy source of food available before winter for free-ranging wisents. Hence, it could be expected that the availability of beech stands within the home range of the wisent population in the Bieszczady Mountains could influence their patterns of habitat use and distribution of their areas of concentration in September–October.
However, our analyses did not confirm a positive influence of availability of beech stands upon the spatial distribution of concentrated areas of wisents during the period of high accessibility of beechnuts. The proportion of beech stands within the 50% kernel areas, identified in the mast years for the September–October period (when the availability of beechnuts was at its maximum), did not exceed its value in the other months of the year and was generally much lower than within the 95% kernel areas, which represents the part of their home range that was actually used.
Therefore, according to our data, the domination of beech stands and the phenomenon of mast years do not play a significant role in habitat selection by wisents belonging to the free-ranging population of the Bieszczady Mountains.

Author Contributions

Conceptualization, K.P. and A.W.-G.; methodology, K.P. and A.W.-G.; software, M.J.; validation, A.W.-G. and M.J.; formal analysis, M.J.; investigation, A.W.-G. and M.J.; resources, A.W.-G. and M.J.; data curation, A.W.-G. and M.J.; writing—original draft preparation, K.P. and A.W.-G.; writing—review and editing, K.P. and A.W.-G.; visualization, M.J.; supervision, K.P.; project administration, A.W.-G. and M.J. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding. Field data were collected during the routine activities of the Carpathian Wildlife Research Station.

Data Availability Statement

The raw data supporting the conclusions of this article will be made available by the authors on reasonable request.

Conflicts of Interest

The authors declare no conflicts of interest.

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Forests 15 01618 g001
Figure 1. Beechnut sampling sites in the Bieszczady Mountains. For every sampled forest district, number of plots in years 2013 and 2022 are given.
Figure 1. Beechnut sampling sites in the Bieszczady Mountains. For every sampled forest district, number of plots in years 2013 and 2022 are given.
Forests 15 01618 g001
Table 1. A comparison of average values of beechnut crop (g dry matter/m2 ± SD) within home ranges of two wisent herds in the Bieszczady Mountains in mast years of 2013 and 2022.
Table 1. A comparison of average values of beechnut crop (g dry matter/m2 ± SD) within home ranges of two wisent herds in the Bieszczady Mountains in mast years of 2013 and 2022.
Year20132022
Herd
Baligród52.8 ± 39.7038.2 ± 30.20
Tworylne107.5 ± 67.1295.4 ± 50.86
Table 2. A comparison of the proportion of beech-dominated tree stands vs. the area of all other types of tree stands within home ranges of two wisent herds of the Bieszczady Mountains between (a) September and October and (b) November and August in the years 2013 and 2022. Except for the Baligród herd in September–October in 2022, in all cases there were statistically significant differences (p ≤ 0.05).
Table 2. A comparison of the proportion of beech-dominated tree stands vs. the area of all other types of tree stands within home ranges of two wisent herds of the Bieszczady Mountains between (a) September and October and (b) November and August in the years 2013 and 2022. Except for the Baligród herd in September–October in 2022, in all cases there were statistically significant differences (p ≤ 0.05).
a.
HerdYear% of Beech Habitats
in 95% Kernel Area		% of Beech Habitats
in 50% Kernel Area
Baligród herd2013386
20224341
Tworylne herd2013264
2022299
b.
HerdYear% of Beech Habitats
in 95% Kernel Area		% of Beech Habitats
in 50% Kernel Area
Baligród herd20134123
20224130
Tworylne herd2013275
2022181
Table 3. A comparison of the occurrence of wisents from two herds of the Bieszczady Mountains during the highest availability of beechnuts (September–October) in years 2013 and 2022: (a) within the whole area of their home ranges, and (b) within forest compartments dominated by beech.
Table 3. A comparison of the occurrence of wisents from two herds of the Bieszczady Mountains during the highest availability of beechnuts (September–October) in years 2013 and 2022: (a) within the whole area of their home ranges, and (b) within forest compartments dominated by beech.
a.
HerdYearNumber within 95% Kernel AreaNumber within 50% Kernel AreaDensity per ha within 95% Kernel AreaDensity per ha within 50% Kernel Area
Baligród herd20135051580.080.48
202211113850.040.18
Tworylne herd20133121380.060.22
20222721010.060.17
b.
HerdYearNumber within 95% Kernel AreaNumber within 50% Kernel AreaDensity per ha within 95% Kernel AreaDensity per ha within 50% Kernel Area
Baligród herd2013134100.060.51
20224291000.040.11
Tworylne herd20134330.030.12
20223080.020.14
Table 4. A comparison of the occurrence of wisents from two herds of the Bieszczady Mountains during months with low or no accessibility of beechnuts (November–August) in years 2012/2013 and 2021/2022: (a) within the whole area of their home ranges and (b) within forest compartments dominated by beech.
Table 4. A comparison of the occurrence of wisents from two herds of the Bieszczady Mountains during months with low or no accessibility of beechnuts (November–August) in years 2012/2013 and 2021/2022: (a) within the whole area of their home ranges and (b) within forest compartments dominated by beech.
a.
HerdYearNumber within 95% Kernel AreaNumber within 50% Kernel AreaDensity per ha within 95% Kernel AreaDensity per ha within 50% Kernel Area
Baligród herd2013345715040.190.55
2022351111010.120.42
Tworylne herd201310,1134761.580.45
2022389911050.804.33
b.
HerdYearNumber within 95% Kernel AreaNumber within 50% Kernel AreaDensity per ha within 95% Kernel AreaDensity per ha within 50% Kernel Area
Baligród herd20136391750.090.28
20228121700.070.21
Tworylne herd2013156170.090.29
202227970.231.99
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Wołoszyn-Gałęza, A.; Januszczak, M.; Perzanowski, K. European Beech Masting Cycles and the Spatial Distribution of Wisents in the Bieszczady Mountains, Poland. Forests 2024, 15, 1618. https://doi.org/10.3390/f15091618

AMA Style

Wołoszyn-Gałęza A, Januszczak M, Perzanowski K. European Beech Masting Cycles and the Spatial Distribution of Wisents in the Bieszczady Mountains, Poland. Forests. 2024; 15(9):1618. https://doi.org/10.3390/f15091618

Chicago/Turabian Style

Wołoszyn-Gałęza, Aleksandra, Maciej Januszczak, and Kajetan Perzanowski. 2024. "European Beech Masting Cycles and the Spatial Distribution of Wisents in the Bieszczady Mountains, Poland" Forests 15, no. 9: 1618. https://doi.org/10.3390/f15091618

APA Style

Wołoszyn-Gałęza, A., Januszczak, M., & Perzanowski, K. (2024). European Beech Masting Cycles and the Spatial Distribution of Wisents in the Bieszczady Mountains, Poland. Forests, 15(9), 1618. https://doi.org/10.3390/f15091618

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