The process of a wolf pack splitting in Białowieża Primeval Forest,Poland

In 1998, the pack of 7 wolvesCanis lupus Linnaeus, 1758, radio-tracked in Białowieża Primeval Forest, East Poland, split into 2 packs (2 and 5 wolves), when an 8-year-old alpha female ceased breeding. The two sister-packs subdivided their original territory, but their ranges overlapped extensively (49%) for one year after the split, except for May-June, when both new packs reared pups. We propose that food related factors could have been the ultimate cause of splitting of a large pack. In European temperate forests, pack size of 5–6 wolves is optimal for the consumption of the red deerCervus elaphus.     

Reproductive behaviour of wild-living wolves in Białowieża Primeval Forest (Poland)

Characteristics of the reproductive behaviour of wolves (Canis lupus) were studied by radio-tracking and snow-tracking of four packs in Białowieża Primeval Forest (BPF), Poland, in 1995–1999. Signs of mating occurred between 12 January and 22 March. Parturition occurred between 19 April and 12 May, and the denning period lasted for 49–64 days. During that time, wolves used 1–3 den sites, spending on average 27 days at each site. The dens were never reused in consecutive years, but year after year the breeding sites were located in the same parts of the pack’s territory. Ten days before parturition pregnant females reduced their normal mobility by half (from 23 to 13.5 km day⁻¹, on average). During the ten days post-partum, they spent 85% of their time with pups and travelled 3.9 km day⁻¹, only. On days 11–30 after parturition, females spent 74% of their time tending pups and increased their daily movement distance to a mean of 13.3 km. The females resumed full mobility 50–70 days after parturition, which coincides with termination of the weaning process. Anecdotal observations indicated that pups were tended by other pack members while the mother was absent. Compared with the years 1947–1950, in 1995–1999 the breeding season of wolves in BPF occurred two weeks earlier. A possible reason was the 1 to 1.5-degree increase in the mean annual temperature during the last 50 years.     

Species composition and activity of bats flying over rivers in the Białowieża Primeval Forest

Bats were netted at two sites over the forest rivers Narewka and Hwoźna in Białowieża Primeval Forest (E Poland), during the summer of 1994 and 1995. A total of 452 bats of 11 species were captured. The number of bats netted each night was positively correlated with the minimum air temperature. The rate at which the three commonest species —Nyctalus noctula, Myotis daubentonii andNyctalus leisleri — were caught varied significantly through the night, with a major peak after sunset. ForN. noctula air temperature was apparently positively related to the size of the morning peak.     

Spatial interactions between grey wolves and Eurasian lynx in Białowieża Primeval Forest, Poland

Various species of large predators are reported to influence each other through interference or exploitation competition that may affect demography and survival of the subordinate species. We analyzed spatial relationships between grey wolf (Canis lupus) and Eurasian lynx (Lynx lynx) in Białowieża Primeval Forest (BPF, eastern Poland) to determine how they partitioned the space. The wolves (n = 8) and lynx (n = 14) were radio-tracked in 1991–1999. Three wolves and seven lynx were radio-tracked simultaneously in 1994–1996. Territories of wolf packs and home ranges of lynx overlapped considerably (76% of wolf territories and 50% of lynx home ranges, on average). In three cases, their core areas were also overlapping. Wolf-lynx dyads with overlapping home ranges were simultaneously located at distances from 0 to 28 km from each other. We found neither avoidance nor attraction between wolves and lynx occupying the same areas. We concluded that in BPF, the two large predators coexist due to specialization on different preferred prey and heterogeneous habitat.     

Soil and vegetation nutrient response to bison carcasses in Białowieża Primeval Forest,Poland

Ungulate carcasses can have important effects on the surrounding soil and vegetation. The impact of six carcasses of European bison (Bison bonasus) was investigated for the first time in a natural temperate forest (Białowieża, Poland) by measuring soil and plant nutrient concentrations along a gradient extending from the centre of each carcass. Calcium concentration and pH were found to be higher at the centre of the carcass, decreasing towards the periphery. This effect lasted up to 7 years after the death of the animal. The concentration of most nutrients in the soil and plants varied irregularly, suggesting an effect of the carcass at its centre but the absence of a clear pattern of variation along the gradient. Concentrations of NO₃⁻ in the soil differed only at the 1-year old carcass, suggesting a fast turnover of nitrate in temperate forests. Our results show that the effects of large herbivore carcasses on soil and plant nutrient concentrations are not easily detectable in a temperate forest as in more homogeneous habitats, such as tundra and prairie. This may be due to the high activity of scavengers and nutrient recycling in the study area, but it may also be a consequence of a more complex and patchy interaction between nutrient availability and other limiting factors in temperate forests.     

Sarcoptic mange vulnerability in carnivores of the Białowieża Primeval Forest,Poland: underlying determinant factors

Sarcoptic mange caused by Sarcoptes scabiei, is a highly contagious worldwide mite infection responsible for epizootic skin disease in populations of wild and domestic mammals. It causes skin disorders that may lead to severe generalized skin disease (mange). We investigated the spread and dynamics of sarcoptic mange in one of the best preserved carnivore communities in the Bia?owie?a Primeval Forest over the last 20 years. Sarcoptic mange was found in 6 of 10 investigated species. The highest percentage of infected individuals was recorded in red foxes (19 %), European badgers (9 %), grey wolves (7 %), raccoon dogs (6 %), and stone martens (5 %). Other species were not infected (least weasel, American mink, European polecat, pine marten) or infected sporadically (Eurasian lynx––1 case). Rate of infection was correlated with the sum of ecological and behavioral features, which may increase the spread of infection. Outbreaks of sarcoptic mange occurred simultaneously in several species of carnivores, which indicates the occurrence of between species transmission. We found that the source of investigated individuals (trapped, culled and found dead) may influence the results of the analysis. Our study shows that in carnivore communities such as in Bia?owie?a Forest, numerous species may serve as a reservoir of the pathogen and favor its persistence in the environment. This study indicates that sociality and use of den sites are important factors affecting sarcoptic mange spread and prevalence in wild populations of carnivores.     

The size and diversity of the soil seed banks and the light requirements of the species in sunny and shady natural communities of the Białowieża Primeval Forest

The research was conducted in two natural forest communities: Potentillo albae-Quercetum (oak forest) which allows much light to reach the forest floor and Tilio-Carpinetum typicum (hornbeam forest) which shades the herb layer heavily. The seed banks were estimated from numbers of seedlings emerging from soil samples over one growing season.(1) Our results confirm the hypothesis that persistent seed banks are mainly formed by species with high light requirements. Of the species found predominantly in the seed bank and absent from the herb layer or occurring there very rarely in both communities 83% of species and 70% of seedlings were strongly light-demanding (Ellenberg's light index 6–9). However, the results do not support the hypothesis that seed banks in natural deciduous forest communities are small, poor in species and do not reflect the species composition of herb layer.(2) The seed banks of both communities were rich in species and relatively large. Species richness in the oak forest turned out to be higher than in the hornbeam forest (51 vs 45 species/2.4 m₂), but size was smaller (2659 vs 5789 seedlings/2.4 m₂). In the oak forest the most abundant species in the seed bank was Galium boreale, but it constituted only 19% of the total number of seedlings, whereas in the hornbearn forest the dominant species, Urtica dioica, constituted 57% of the total.(3) In each community the species composition of the seed bank and the herb layer was very similar (>70%).(4) The seed bank was more diverse in the oak forest than in the hornbeam forest (H 2.34 vs 1.68).(5) The seed banks of both communities differed in the contribution of species with varied light requirements; in the sunny oak forest species with high light requirements dominated, whereas in the shady hornbeam forest both strongly and moderately light-demanding species had similar contributions.Nomenclature: Follows Ehrendorfer (1973) and Matuszkiewicz (1981).     

The mosaic diversity of field layer vegetation in the natural and exploited forests of Białowieża

The main focus of the study was to find out whether and how the forestry management in the Biaowiea Forest has influenced the mosaic diversity of field layer vegetation and habitat heterogeneity in the temperate hard wood forest (Tilio-Carpinetum). Five belt-transects represent natural and semi-natural old-growths in Biaowiea National Park (BNP), the commercial forest (CF), and secondary stands in the CF. Along the transects, distinct patches of field layer vegetation were identified and delimited, and their species composition was assessed. Along central lines of the transects the environmental data was collected from sample points established at regular intervals: soil pH and the distances to the 4 closest trees, their species, and dbh (diameter at the breast height=135 cm). The group of patches (on average 50 per transect) was subjected to the dichotomous classification in order to identify a number of characteristic types of microphytocoenoses – synusiae. Beta-diversity of the vegetation was expressed with the Simpson diversity index calculated on the basis of areas occupied by synusiae. The most structurally diversified were the transects representing natural old-growths in the BNP. A calculation of similarity coefficients between the transects on the basis of their mosaic composition displayed sharper differences between old-growth and secondary communities in summer than those found in spring. Forestry management has seriously modified the tree species composition in two studied stands, through planting oak or initiating succession. This resulted, both in plantation and pioneer stand, in loss of the main components of natural old-growths: hornbeam (Carpinus betulus), Norway spruce (Picea abies), lime (Tilia cordata) and maple (Acer platanoides). Significantly a wider range of soil pH values in natural old-growths coincided with a higher diversity of herbaceous mosaics comparing to secondary stands. The elimination of spruce (the species being most commonly blown down, projecting an acid canopy shadow) in two studied areas, elimination of dead wood, and destruction of humus layers through ploughing, could contribute to a reduction of specific microenvironments. A lower mosaic diversity of secondary stands coinciding with the unification of environmental conditions along the transects may indicate that forestry management can lead to the disappearance of a highly structural character in natural forest communities.Nomenclature: Flora europaea; Sokoowski (1993).     

Resting site selection by large herbivores – The case of European bison (Bison bonasus) in Białowieża Primeval Forest

Resting site selection by European bison (Bison bonasus, L., 1758), the largest terrestrial mammal of Europe, was studied in the free-ranging population in Bia?owie?a Primeval Forest (Poland) in 2009–2010. In total, 104 sites of 21 bison (both collared and uncollared) were analysed to determine the most important microhabitat characteristics selected by resting bison during summer and winter and to study the influence of supplementary feeding on resting behaviour of this herbivore. Resting sites were identified on the basis of GPS locations and activity records collected by GPS collars, as well as direct observations of bison, and were compared with control sites. Microhabitat selection by bison did not differ significantly between the sexes. During summer and winter, bison resting sites displayed a high tree density, low visibility and high complexity (structures providing cover). Summer resting sites were also characterised by a significantly lower abundance of blood-sucking insects and denser canopy than control sites. Winter resting sites showed a lower complexity and higher visibility than summer sites, and were less often located in mixed forest habitats. During winter, bison rested more frequently in forest below 50 years of age than in older forest. Resting sites of non-fed bison were more often located in young coniferous forests, were lower in visibility and situated closer to open areas than sites of bison using supplementary winter feeding, suggesting a trade-off between food and cover. The results indicate that European bison select their resting sites in areas of mosaic habitat structure providing cover from disturbances with access to profitable natural forage grounds.     

Seasonal and sexual variation in diel activity rhythms of pine martenMartes martes in the Białowieża National Park (Poland)

From 1991–1996, the activity rhythms of 14 radio-collared pine martensMartes martes (Linnaeus, 1758) (6 males and 8 females) were studied in the pristine deciduous and mixed forests of the Białowieża National Park. Tracking data (5823 h) indicated that the activity rhythms of pine martens varied between sexes and seasons. In spring, male activity peaked at 20.00–00.00 h, whereas in summer and autumn-winter, activity was bimodal, peaking at 18.00–22.00 h and 02.00–04.00 h. Female activity in spring was more evenly distributed than that of males, but in summer their activity peaked at 20.00–00.00 h, while in autumn-winter females had a bimodal rhythm with peaks at 18.00–20.00 h and 02.00–06.00 h. In breeding females, activity rhythms changed in the course of pregnancy and nursing. On average, martens started their activity 73±209 (SD) min before sunset and finished 87±245 min after sunrise. Females became active earlier than males but both sexes terminated activity at the same time. For both males and females the daily activity rhythm was not related to the diurnal course of temperature.     

Lichenicolous fungi are more specialized than their lichen hosts in primeval forest ecosystems,Białowieża Forest,northeast Poland

Taxonomy and diversity of symptomatic lichenicolous fungi (visible as fruitbodies on lichen thalli, their discoloration, and/or deformation) and their specificity to lichen hosts is becoming more and more studied. However, information on their ecology is still scarce. We assess how large the specialization of these fungi towards their hosts and microhabitat is. Epiphytic, epixylic and epigeic lichens and associated lichenicolous fungi were studied on 144 permanent plots in Białowieża Forest in relation to forest communities, species of tree phorophyte and substrates. On all these three studied levels lichenicolous fungi were more specialized than their lichen hosts. Our study provides the first estimation of ecological dependences between associations of lichenicolous fungi and their hosts, microhabitats and forest communities in a primeval forest ecosystem representative of lowland Europe.     

Diversity of the Scuttle Fly (Diptera: Phoridae) Communities in the Plantations of Moist Pine Forests of the Białowieża Primeval Forest and the Tuchola Forest (Poland)

Scuttle fly communities of pine plantations were investigated in 1986 and 1987 in two sites, Bia?owie?a Primeval Forest and Tuchola Forest. Flies were collected using yellow plastic bowls placed on the ground. Up to now 108 species were identifined, and 46 occurred on both sites. Although the number of species was very similar in Tuchola Forest (77) and Bia?owie?a Primeval Forest (75) species diversity was considerably lower on the former site (p<0.05). Ten dominant species were common: Megaselia brevicostalis, M. giraudii, M. manicata, M. nigriceps, M. pleuralis, M. pulicaria-complex, M. pumila, M. verralli, Metopina oligoneura and Triphleba opaca. Five of these were characteristic of both communities (Megaselia verrali , M. brevicostalis, M. pumila, Metopina oligoneura and Triphleba opaca). Similarity of qualitative composition for dominants was rather high (Sø = 0.67), but the quantitative similarity was low (Mo = 0.26). During two study seasons in the community of the Bia?owie?a Primeval Forest the dominance structure did not change markedly and Megaselia verralli was the dominant (over 20%). M. verralli was also the dominant (ca. 30%) in the communities in the Tuchola Forest in 1986, but next year M. pulicaria-complex (ca.50%) dominated. Most of the dominant species are multivoltines and generalists. In Tuchola Forest the disturbances caused by the chemical treatment against Neodiprion sertifer might be the main factor changing the dominance structure in the phorid community in 1987 (extremely high dominance of Megaselia pulicaria-complex).     

The regressive Cirsium rivulare population in an unmown meadow in Białowieża National Park: Influenced by other clonal plants and necromass

This study deals with population dynamics of Cirsium rivulare (brook thistle) present in a meadow that undergoes succession as a result of former disuse. The effect of the following in the succession, currently dominant clonal species and necromass on the performance of the brook thistle population (presence, size of clusters and reproduction) was investigated. The regression stage, the Cirsium rivulare population had attained after mowing, was discontinued. Different kinds of influence from the contiguous species on the brook thistle population could be identified. In the case of Filipendula ulmaria and Carex cespitosa, Cirsium rivulare clusters were found to be present on the edge of the dominant patches. In case of Carex acutiformis patches brook thistle ramets did not form clusters but were present among the sedge ramets. No Cirsium rivulare ramets were found in Phragmites australis and willow (Salix cinerea, S. pentandra and S. aurita) patches. The highest number of generative brook thistle ramets were observed in a Carex acutiformis patch, whereas the highest number of inflorescences on the shoots of the analysed species were identified in the patch dominated by Filipendula ulmaria.Experimental vegetation gaps were created in patches dominated by various species to determine the effect of necromass on the presence of Cirsium rivulare seedlings. A considerable rise in the number of Cirsium rivulare seedlings was noted in Carex acutiformis and Filipendula ulmaria patches following necromass removal in comparison with plots containing necromass. On the other hand, removing the necromass and foliage did not contribute to an increase in the numbers of brook thistle seedlings in the patch dominated by Cirsium rivulare itself.     

Movements of European bison (Bison bonasus) beyond the Białowieża Forest (NE Poland): range expansion or partial migrations?

The European bison is the largest terrestrial mammal in Europe. After extinction in the wild at the beginning of the twentieth century, it was re-introduced to Bia?owie?a Forest and other woodlands in Eastern Europe. In this paper, we analysed the movements of European bison beyond the continuous woodlands of the Bia?owie?a Forest (NE Poland) between 1964 and 2010. In total, 1,117 direct observations of bison were collected. The number of males moving out of the forest fluctuated during the study period, whilst the number of females steadily increased. The number of male observations outside of the forest per annum was dependent on the population size and snow depth, whilst the number of cows in mixed groups moving outside of the forest was correlated with the population size only. Males were observed mainly alone (50 % of observations) or in small groups of two to three individuals (25 %); however, distribution of group size differed from those observed in the population. There was a significant difference between the direction of movement of males and females out of the forest—males moved mainly west and southwest, whilst females moved to the north. This was also significantly different from the expected movement direction. The mean distance of bison observations from the forest border was 1.8?±?0.13 km and did not differ significantly between sexes. After 1990, males were observed significantly farther away from the forest (2.2 km) than in the previous years (0.9 km). Most observations (94 % of bulls and 93 % of cows) were up to 5 km from the forest edge. The range of bison in the vicinity of the Bia?owie?a Forest was strictly seasonal. Most observations (78 % in males and 88 % in females) were recorded from November to April. Increasing utilisation of areas beyond the forest habitats may be driven by different factors but most probably it is related to range expansion and the bison's preference for open habitats. The strong seasonal pattern of bison movements indicate that the partial seasonal migrations were initiated in the Bia?owie?a population.     

Factors affecting daily ranges of red deer<Emphasis Type="Italic">Cervus elaphus</Emphasis> in Białowieża Primeval Forest,Poland

Daily ranges of 19 (6 males, 13 females) adult red deerCervus elaphus Linnaeus, 1758 were studied using 24-h tracking sessions in Białowieża Primeval Forest (BPF), Poland, from 2001 to 2004. Overall, size of mean (± SE) daily ranges was larger for males (1.22 ± 0.10 km²) than females (1.00 ± 0.09 km²), although the difference was not significant. Similarly, mean daily ranges were 6–46% larger for males than females in each season, although there were no statistical differences in mean daily ranges among seasons for each sex. Abiotic factors, especially temperature, significantly affected daily ranges of females, but not males, suggesting sexual differences in response to weather variables. On a daily basis, males used 3% of their annual home range, whereas females used 12% of their annual home range, indicating females used their annual home ranges more intensely than males. Consecutive daily ranges overlapped little for each sex. Daily ranges of red deer in BPF were considerably larger than previously reported in Europe, suggesting factors unique to BPF also influenced size of daily ranges.     

Seasonal and spatial pattern of shelter use by badgers<Emphasis Type="Italic">Meles meles</Emphasis> in Białowieża Primeval Forest (Poland)

In 1997–2001, we investigated the use of day-time shelters by radio-collared badgersMeles meles (Linnaeus, 1758) in the Białowieża Primeval Forest, eastern Poland. Each social group of badgers utilised, on average, 9 different shelters per territory (range: 4–20). The main setts, occupied for breeding and winter sleep, were also most frequently used for day-time rest throughout the year (73% of days). Badgers living in the pristine oldgrowth stands utilised larger number of shelters and spent more days in hollow trees (mainly limeTilia cordata), compared to badgers inhabiting younger secondary tree stands. Number of shelters used by individuals varied between seasons and depended on sex and age of animals. In summer, badgers used more shelters than in spring and autumn. In winter, they stayed in their main setts only. Adult males occupied more shelters and spent fewer days in the main sett than other badgers. In spring, females rearing young used only the main setts. The average underground space used by badgers within the main sett was 128 m². It was largest in summer and smallest in winter, and also varied between males and females. We proposed that, in a low-density population, badgers used several setts and other daily shelters to reduce energy expenditure when exploring their large territories and foraging. Furthermore, setts may play a role of marking sites. Analysis of the biogeographical pattern of sett use by European badgers showed that the number of setts used by social groups increased with increasing territory size, whereas the density of setts (n setts/km²) was negatively correlated with territory size. We proposed that different factors could shape the utilisation of setts by badgers in low- and high-density populations.     

Rapid morphological divergence in two closely related and co-occurring species over the last 50 years

We studied morphological variation in two closely related and ecologically similar species of mice of the genus Peromyscus, the deer mouse (P. maniculatus) and white-footed mouse (P. leucopus), over the last 50 years in Southern Quebec. We found that contemporary populations of the two species are distinct in morphology and interpret this differentiation as a reflection of resource partitioning, a mechanism favouring their local coexistence. While there was no size trend, geographic or temporal, both species displayed a concomitant change in the shape of their skull over the last 50 years, although this change was much more apparent in the white-footed mouse. As a result, the two species diverged over time and became more distinct in their morphology. The observed changes in morphology are large given the short time scale. During this period, there was also a shift in abundance of the two species in Southern Quebec, consistent with the northern displacement of the range of the white-footed mouse in the last 15 years. Our study thus reports the changes in morphology of two co-occurring mammal species that were accompanied by changes in distribution and local abundance, potentially in response to rapid climate change.     

Discordant patterns of morphological and genetic divergence in the closely related species Schizanthus hookeri and S.?grahamii (Solanaceae)

In this study I have examined the patterns of morphological and genetic differentiation between two species of the Andean genus Schizanthus that differ in their pollination and mating systems. Schizanthus hookeri has a bee pollination syndrome and is strongly dependent on pollinators for seed set. In contrast, S.?grahamii has a hummingbird pollination syndrome and exhibits late autonomous selfing. Southern populations of the latter species have red flowers (reddish morph), while northern populations have yellow (yellowish morph) or pink flowers (pinkish morph). I used two noncoding chloroplast DNA (cpDNA) regions to investigate the genetic affinities between S.?hookeri and the three morphs of S.?grahamii. I also performed intra- and interspecific crosses to assess whether gene flow between species was possible. Phylogenetic analyses supported the existence of two differentiated clades that did not match currently accepted taxonomic classification. Accordingly, genetic distance did not correlate significantly with morphological distance. No fruits were produced from interspecific crosses, and there were no individuals with intermediate morphology that could indicate current and frequent hybridization events between species.?I propose that the discordance between cpDNA data and conventional taxonomy could be explained by parallel evolution, or alternatively, by a very sporadic hybridization.