Leśne Prace Badawcze / Forest Research Papers
Grudzień / December 2016, Vol. 77 (4): 302–323
Available online: www.lesne-prace-badawcze.pl
Submitted: 1.08.2016, reviewed: 3.08.2016, accepted after revision: 12.08.2016
© 2016 A. Kujawa et al.
The Białowieża Forest – a UNESCO Natural Heritage Site – protection priorities
Anna Kujawa1*, Anna Orczewska2, Michał Falkowski3, Malgorzata Blicharska4, Adam Bohdan5, Lech Buchholz6,
Przemysław Chylarecki7, Jerzy M. Gutowski8, Małgorzata Latałowa9, Robert W. Mysłajek10, Sabina Nowak11,
Wiesław Walankiewicz12, Anna Zalewska13
Institute for Agricultural and Forest Environment, Polish Academy of Sciences, Bukowska 19, 60–809 Poznań, Poland;
Department of Ecology,
Faculty of Biology and Environmental Protection, University of Silesia, Bankowa 9, 40–007 Katowice, Poland;
Ornithological Society, Radomska 7, 26-760, Pionki, Poland;
Uppsala University, Department of Earth Sciences, Villavägen 16, 75 236 Uppsala,
Foundation “Dzika Polska”, Petoego 7 lok. 18, 01–917 Warszawa, Poland;
Polish Entomological Society, Dąbrowskiego 159, 60–594
Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00–679 Warszawa, Poland;
Forest Research Institute,
Department of Natural Forests, Park Dyrekcyjny 6, 17–230 Białowieża, Poland;
University of Gdańsk, Department of Plant Ecology, Laboratory
of Paleoecology and Archaeobotany, Wita Stwosza 59, 80–308 Gdańsk, Poland;
University of Warsaw, Faculty of Biology, Institute of Genetics
and Biotechnology, Pawińskiego 5a, 02–106 Warszawa, Poland;
Association for Nature “Wolf”, Twardorzeczka 229, 34–324 Lipowa, Poland,
Institute of Biology, Siedlce University of Natural Sciences and Humanities Prusa 12, 08–110 Siedlce, Poland;
Warmia and-Mazury University
in Olsztyn, Faculty of Biology and Biotechnology, Department of Botany and Nature Protection, pl. Łódzki 1, 10–727 Olsztyn, Poland
*Tel. +48 65 5134253, e-mail: firstname.lastname@example.org
Abstract. Despite the fact that only parts of the Białowieża Forest are protected as a national park and nature reserves, the forest
is nevertheless as a whole considered a UNESCO Natural Heritage Site, Biosphere Reserve and an integrated Natura 2000 site.
In the presently ongoing debate on the conservation priorities regarding the natural value of this forest and the current bark beetle
outbreak, two distinct approaches can be recognized: (1) management assumed to involve considerable interference with the forest
ecosystems; (2) maintenance of ecological processes and spontaneous restoration of the forest communities. The Białowieża
Forest – especially its strictly protected parts – is a “bastion” where species characteristic of ancient forests (including so-called
primeval forest relicts) have survived until today. This has been achieved by maintaining the forest’s complexity in areas with
considerably reduced human inuence, but most of all by maintaining a full spectrum of forest communities, naturally developing
forests diverse in age, species composition and spatial structure including stand dieback and breakdown. The following factors
need to be taken into account in the protection of the Natural Heritage Site: (1) the internationally recognized value of the
Białowieża Forest including its biodiversity, the level of preservation of forest communities and the ongoing natural processes;
(2) existing documents and policies concerning nature conservation; (3) research ndings from the Białowieża Forest and other
natural forest complexes. The key priority is to limit any activities in this forest to an indispensable minimum, mostly concerning
security close to roads and tourist tracks as well as collection of re wood by locals. Without this strict protection, successive
and slow anthropogenic transformation will result in the Białowieża Forest sharing the same fate as other forest complexes of the
temperate climate zone in Europe or America and lose its globally appreciated value.
Keywords: natural forests, nature protection, biodiversity, natural ecological processes, forest management
The Białowieża Forest (BF) includes a coherent forest
complex, located on the border between Poland and Be-
larus. The whole Belarusian part is under legal protection
as a national park. The Forest’s part on the Polish side
comprises an area of 62,000 ha: the Białowieża Natio-
nal Park (BNP), 10,500 ha; a network of nature reserves,
12,000 ha; and managed forests, 39,500 ha (Wesołowski
et al. 2016).
303A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
The uniqueness of BF nature, outstanding natural values for
biodiversity conservation, representative exemplication of on-
going ecological and biological processes of great importance
in ecosystem evolution and progress, as well as scientic val-
ues beyond measure have been recognised on the international
forum. The BF was inscribed on the United Nations Education-
al, Scientic and Cultural Organisation (UNESCO) World Her-
itage List, as the rst transboundary site (Polish–Belarusian) in
Europe (on the Polish side, initially just the BNP was included
in the List). In 2014, at the joint Nomination Dossier of Po-
land and Belarus, the whole area of the Forest was nominated
the Natural World Heritage site, based on the evaluation crite-
ria (ix) and (x). In Poland, the entry borders do not include the
Forest’s edges in the immediate vicinity of the village Hajnów-
ka and villages situated along the western border of the Forest
(Polski Komitet ds. UNESCO 2016a).
The BF is a member of the UNESCO World Network of
Biosphere Reserves (WNBR) of the Program Man and Bios-
phere (MAB). According to Article 4 (Criteria) of the WNBR
Statutory Framework, a designated biosphere reserve should
encompass a mosaic of ecological systems that represents the
main biogeographic regions in a given country. The reserve is
designated with the aim to establish the sites for nature con-
servation, observation and research. Each reserve consists of
three zones: the core zone, the buffer zone and the transition
zone (Battisse 1982; UNESCO 1984; Denisiuk, Witkowski
1990; Polski Komitet ds. UNESCO 2016b). When established
in 1976, the Biosphere Reserve Białowieża encompassed just
the area of the BNP. Since 2005, the Reserve has embraced
the areas of the Forest Districts: Białowieża, Browsk and Haj-
nówka, as well as the areas of the following municipalities (in
part or in whole): Białowieża, Hajnówka, Dubicze Cerkiewne,
Narew, Kleszczele, Narewka and Czeremcha. At the same time,
the BF is protected as the Special Protection Area PLC200004
under the Natura 2000 Network (Zarządzenie 2011), the Impor-
tant Bird Area (IBA PL046) (BirdLife International 2016) as
well as the Area of Protected Landscape (almost entire BF area)
(Rozporządzenie 2005) and the Forest Promotional Complex
‘Białowieża Forest’ (excluding BNP) (Zarządzenie 1994).
At the turn of 2016, in Poland, long-standing disputes
between the supporters of implementation of current forest
management practices in the managed parts of the BF1 and
advocates of giving the priority to the protection of natural
processes in forest ecosystems reemerged (Wesołowski et
al. 2016). Actually, the conict over the scope and method
of nature conservation in the BF has been lasting for years
whilst it intensied for the rst time in the early 1990s (Szu-
jecki 2008; Blicharska, Angelstam 2010). The essence of the
dispute is the disagreement over how the forest should be per-
ceived and how the BF’s natural values should be evaluated
and protected now and in the future. The foresters responsible
for the forest management in the managed part of the Forest
and some scientists associated with them claim that human in-
tervention is necessary to assure the survival of the BF. On the
other hand, the majority of scientists (mainly biologists and
also foresters) and numerous non-governmental organisations
postulate allowing the BF ecosystems for spontaneous deve-
lopment, keeping the level of human intervention at minimum
(amongst others, Blicharska, Van Herzele 2015; Blicharska et
al. 2016; Wesołowski et al. 2016).
The aim of the paper is to demonstrate the natural values
of the BF and to determine its protection priorities, based
on the scientic data and in the light of the mandatory legal
documents and acts regarding the protection of this area.
2. The history of the Białowieża Forest
Today’s woodland of the BF is a result of ecological pro-
cesses, shaping vegetation already from the beginning of the
Holocene that is, for 12,000 years. The course of subsequent
stages of vegetation development followed the pattern cha-
racteristic for northern regions of east-central Europe, which
in the early Holocene comprised of pioneer pine-birch forests
and later elm and hazel expansion (about 11, 300 and 10, 500
years ago, respectively), followed by the development of mul-
ti-species deciduous forests 9,300–3,800 years ago and then
formation of multi-layer deciduous forest (Carpinion betuli)
with dominating hornbeams (about 3,800 years ago). Climate
was the main factor initiating particular stages of changes in
plant communities, whilst spruce expansion that occurred re-
latively late in the region (1,500 years ago) was most probably
a combined effect of earlier human impact in the Roman Pe-
riod and climate changes (Zimny 2014; Latałowa et al. 2016).
Palynology allows for studying the long-term dynamics of
forest communities and provides information for not only ap-
praisal of the current ecological processes in view of the proper,
long-term perspective but also prediction of possible scenarios
of future vegetation changes under the conditions of changing
climate. In recent times, declining groundwater levels have
been the most important factor leading to the transformation
of almost all habitats in the BF (Pierzgalski et al. 2002). The
process of spruce decline caused by the European spruce bark
beetle invasion under the conditions of recurring long-term
droughts represents the most spectacular example of that (Ke-
czyński 2002). Palynological data indicate that during the last
millennium, spruce population uctuated in short- and long-
term periods of time (Latałowa et al. 2016). Ecological me-
chanisms of growth or reduction of spruce population within
various habitats, because of a range of forest management me-
thods, ground res, excessive herbivore pressure and declining
groundwater levels in hydrogenic habitats have been discussed
by many authors (e.g. Faliński 1986; Keczyński 2005, 2007;
Czerepko 2008; Kuijper et al. 2010; Niklasson et al. 2010;
304 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
Bobiec 2012; Bobiec, Bobiec 2012). It can also be assumed
that at least some of the recurring radical reductions of spruce
populations in the past, as reected by pollen data, have been
due to European spruce bark beetle outbreaks (Latałowa et al.
2016). The history of spruce in the BF shows that the species
has endured in good condition both the periods of drought and
unfavourable effects of other ecological factors and is still one
of the main forests forming tree species in the area. Such high
capability of population restitution was probably related to the
broad phytocoenotic range of spruce in this region (Faliński
1986; Sokołowski 2004), which should be recognised as the
positive aspect when assessing the potential ability of the spe-
cies to survive the current and future outbreaks of the European
spruce bark beetle.
The BF did not avoid economic exploitation in the past (Fa-
liński 1986; Samojlik 2010; Samojlik et al. 2013); nonethe-
less, the assessments based on palynological indices of human
activities clearly indicate that an extent of human-induced
transformations has been relatively low, both in the prehistory
and at the historical times (Latałowa et al. 2015, 2016). This
has been possible because of an underdeveloped settlement
network, nearly in all the archaeological periods (Wawrusie-
wicz 2011; Jaskanis 2012), as well as in medieval and modern
times (Mikusińska et al. 2013; Samojlik et al. 2013). The spe-
cial status of a royal asset limited forest exploitation and agri-
cultural expansion in the BF from the 15th to the beginning
of the 20th century (Samojlik et al. 2013). The comparison
of pollen shares of both cultivated plants and those charac-
teristic for habitats changed because of human activities re-
corded in pollen diagrams from the BF sites and from other
sites in northern Poland (Latałowa et al. 2016) shows a great
uniqueness of the BF material. The latter is characterised by
not only a considerably lower proportion of plant taxa typical
for anthropogenic vegetation but also a diminutive incidence
of agricultural indices. This explains the exceptional and to a
high degree natural present state of forest communities in the
BF. Harvesting and processing forest raw materials as well
as grazing were the main forms of the BF utilization in the
past (Hedemann 1939; Samojlik 2010; Samojlik et al. 2016),
which allowed for maintaining continuity of forest habitats
along with the successful natural regeneration processes. This
contributed to the maintenance of presently best preserved
fragments of deciduous and mixed forests that occur in the
North European Plain, and that is why the BF has been nomi-
nated the Natural World Heritage site by UNESCO.
3. Nature conservation in the Białowieża Forest
– present status and contemporary threats
The nature of the BF is protected under mandatory legal re-
gulations regarding conservation of animal, plant and fungi spe-
cies, and conservation plans comprising conservation tasks for
the BNP, the nature reserves as well as the habitats and species
protected under the ecological network Natura 2000 encompas-
sing the entire area of the Forest. Furthermore, the recommen-
dations and nature conservation priorities are accentuated in the
management plan for the Forest Promotional Complex ‘Puszcza
Białowieska’ for the years 2012–2021 (RDLP 2011) as well as
in the Nomination Dossier to the UNESCO for the inscription
of the BF on the World Heritage List (Krzyściak-Kosińska et
al. 2012). Various informal proposals were also included in the
package of projects of legislation acts on designation of the na-
tional park within the entire area of the BF. The latter was ela-
borated by the team appointed by the President of Poland for the
duration of the works on the legal act aiming at the regulation
of the status of national natural and cultural heritage of the BF
(Projekty ustaw 2006). It should be stressed that the BF was
inscribed on the List of UNESCO World Heritage Sites, based
on the selection criteria that clearly determine the priorities –
the protection of ongoing ecological and biological processes in
natural habitats representative for in-situ conservation of biolo-
gical diversity (Polski Komitet ds. UNESCO 2016a).
Even though the BF is hardly a primeval forest (never di-
sturbed by humans), its ecosystems are best preserved in the
European Lowlands. For that reason, spontaneous ecologi-
cal processes ongoing here – as a consequence of numerous
factors at the global, regional and local levels (i.e. the climate
change, changes in water regime, nitrogen deposition, and uc-
tuations in large herbivore populations, as well as a cessation
of livestock grazing in the managed part of the BF) – should
be recognised as the a priority over economic forest functions.
The frequently heard arguments about allegedly harmful effects
of long-lasting (more than 90 years) strict protection on forest
biotic diversity in the BNP (Brzeziecki et al. 2016) have three
essential weaknesses. First of all, as stressed by Jaroszewicz
et al. (2016), such conclusions are based on the observations
carried out on an area of only 15.4 ha, in various types of forest
communities, with diverse hydrological and edaphic conditions
where the dynamics of the woody species populations operate
at different scales. Hence, because of their spatial and temporal
scale, the demographic processes observed by Brzeziecki et al.
(2016) are not representative enough for the BF as a whole.
Second, the latter study neglects the importance and absolute
uniqueness of an opportunity to register the course of natural
ecological processes under the conditions uninterrupted by fo-
rest management activities, which is assured exclusively by the
strict protection of forest ecosystems in the Park. The observed
directions and rates of natural ecological processes should not
be the subject of evaluation. Hence, the statements that long-
term strict protection has adverse effects on nature since it leads
to biodiversity loss (Brzeziecki 2016) are not justied. In the
ecological perspective, the recently observed decrease in the
abundance of some tree species (e.g. oak, spruce, aspen, birch,
305A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
pine, and ash) can be the combined effect of different natural
factors, such as increased herbivore pressure and eutrophica-
tion, global warming, groundwater levels decline, pathogenic
infections and, nally, the effect of secondary succession. The-
refore, this decrease is not a result of the strict protection in
the BNP. The response of individual plant species is a result of
competition, which under the inuence of certain factors, leads
to favouring some species at the cost of others. The possibili-
ty to investigate these processes is intrinsically valuable, and
for such studies, the BF is the most appropriate site within the
North European Lowlands. The statements presented in scien-
tic literature, which emphasise the great importance of forest
management for the preservation of biotic diversity (Brzeziecki
et al. 2016), express lack of understanding or the ignorance of
the essential value of the BF, which is referred to in the UNE-
SCO criterion IX. This criterion points out high naturalness of
forests and the obligation to protect natural processes that sho-
uld be accompanied by the minimisation of human interference.
Furthermore, as Weiner (2016) writes, the aforesaid statements
are ‘a form of contestation of the view that the existence of a
natural forest ecosystem is a priceless value, worth preserving’
and the manifestation of ‘a serious crisis of values’. The third
weakness of the above-mentioned argumentation as to alleged
detrimental effects of the strict protection is the identication
of the observed demographic processes in tree species growing
in the strict nature reserve, that is, decrease in the population
numbers of certain species followed by the expansion of others,
mainly hornbeam and lime, as the evidence of the homogenisa-
tion of the Forest’s plant communities (Drozdowski et al. 2012;
Brzeziecki et al. 2016). In the recent years, the term ‘homogeni-
sation’ has been repeatedly evoked (e.g. McKinney, Lockwood
1999; Olden et al. 2004; Naaf, Wulf 2010). Nonetheless, the
results of studies carried out in the BF managed forests (Droz-
dowski et al. 2012), referred to by Jaroszewicz et al. (2016),
evidently show the same dynamic trends regarding the tree spe-
cies that are observed in the protected forests in the nature re-
serve. Drozdowski et al. (2012) state that ‘the process of forest
community homogenization in old-growth forests in the mana-
ged parts of BF has not so far been as advanced as that in the
BNP Strict Reserve’. As emphasised by the authors themselves,
this is due to ‘the forest management activities (regeneration,
stock tending operations, stand structure regulation), which
improved the growth conditions of the threatened tree species.
Therefore, the maintenance of high tree species richness, which
assures safeguarding high natural values of the managed forest
stands was promoted’. Thus, as maintained by Drozdowski et
al. (2012), the higher tree species richness in the managed forest
stands compared to the forests released from human pressure is
attributable to the effects of silvicultural treatments. In contrast,
these processes should be rather explained by the intermedia-
te disturbance hypothesis (IDH), described by Connell (1978)
and widely recognised in ecology. Consistent with IDH, distur-
bances at an intermediate level maximise species richness. The
latter decreases at both low and high level of disturbance. In the
case of the BF, an increase of species richness is of no value in
itself, because such an increase is often connected with the oc-
currence of the light-demanding species, typical for early stages
of forest development (e.g. birch and common aspen), which
in mature forests are successively replaced by shade-tolerant
tree species. At the same time, disturbances can contribute to
the penetration of a forest by alien species, including the inva-
sive ones (Catford et al. 2012), which are a hazard to the entire
forest complex. In view of the fact that disturbances, including
forest management activities, are responsible for the increase in
the species richness, often because of the temporal appearance
of ephemeral species in the forest groundcover, lower species
richness in forests free of human intervention is regularly ob-
served when compared to forests strongly affected by mana-
gement activities. Consequently, the total number of species is
not sufciently objective, or at least not the only one indicator
of a forest conservation value because it often reects the pre-
sence of disturbances affecting the local environment (Boch et
al. 2013). It is worth stressing that the above-described effect is
often observed in fragmented forests of small area. However,
when considering the larger scale (landscape), it appears that
when compared to managed forests, the number of species in
protected forests is not necessarily lower. In protected forests,
abundance of species populations is often lower, but the total
species richness is higher. Unfortunately, empirical data on
these aspects are almost non-existing, because of the fact that in
Europe, besides the BF, there is a lack of extensive, large forest
complexes, undisturbed by management practices that could be
used in relevant comparative studies.
4. Evaluation of the conservation status of
Natura 2000 sites designated within the area
of the Białowieża Forest and recommendations
on their protection
In accordance with the Standard Data Form (SDF) for the
site Natura 2000 Puszcza Białowieska (BF), 10 habitats are
protected here (5 non-forest and 5 forest habitats) (Table.1). Fo-
rest habitats encompass 67.53% of the area of the Natura 2000
site (63, 147.58 ha) that was designated in the complex with fo-
rest plant communities characteristic of primeval plant associa-
tions (Matuszkiewicz 2007a), much better preserved and more
stable compared to other regions. In the area, deciduous forests
(9170 Galio-Carpinetum, Tilio-Carpinetum oak-hornbeam fo-
rests), which cover 39, 814.56 ha (91.54% of the total area of
Natura 2000 habitats), prevail. The conservation status of this
habitat is assessed as excellent, based on the parameters such
as ‘specic structure and functions’ (serves to dene the typi-
306 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
cal nature of habitat development and conformity with specic
species composition) as well as ‘habitat conservation prospects’
(SDF 2014). Similar overall assessments are reported for ha-
bitats 91E0 and 91F0 (riparian forests) (Table 1). However, it
must be emphasised that in the case of deciduous forests, plant
communities in the BNP or the nature reserves considerably
differ from those of managed forests. In the latter, high diversity
of actual vegetation is observed, which is associated with plant
species occurring after tree harvesting and different stages of
forest regeneration (Kwiatkowski 1994). Managed forests con-
siderably differ from the forests of BNP in terms of stand age
structure, proportions of deciduous species, amount of dead-
wood and biomass. The protected and managed deciduous fo-
rests also differ in terms of biogeochemical cycles. In managed
deciduous forests, the cycle is disturbed as a result of changes
in species composition (e.g. spruce domination). Continuous
losses of mineral nutrients caused by timber removal are ob-
served; water retention is altered, and surface runoff is enhan-
ced within harvesting areas (Kwiatkowski 1994). According
to Matuszkiewicz (2007b), the majority of tree stands within
the BNP entered the decomposition stage, and this instigates
natural uctuations in deciduous forest, that is, the processes
of regeneration ongoing within small areas connect with those
of degeneration linked to tree falls because of natural factors.
Slow changes in the Park’s stands are associated with an incre-
asing dominance of hornbeam and lime trees and the reduction
in the shares of other tree species (especially spruce, birch and
aspen) (Matuszkiewicz 2007b).
In line with the conservation action plan for the Natura
2000 site PLC200004 Puszcza Białowieska (Zarządzenie
2015), the conservation tasks rule out management activities
in forest habitats 91D0 and 91E0. In the case of habitat 9170,
all the stands with more than 10% share of trees older than 100
years are to be excluded from forest management activities
(Table 2). It needs to be highlighted that the only conservation
task proposed for the non-forest habitats (except for 3150) is
improvement in knowledge through inventories aiming at re-
cognition of all the habitat sites and monitoring of their con-
servation status with the use of methods established by the
Chief Inspectorate of Environmental Protection, Poland.
Forest management is classied as a potential threat
(Table 2) and not as an existing threat, and this is probably
due to the standpoint of the Minister of Environment (do-
cument DP-074-60/30110/15/JJ of 13 August, 2015), which
states that the fundamental method of conservation of Natu-
ra 2000 habitats is sustainable management of nature reso-
urces. As only unsustainable forest management threatens
protected habitats, and the principles of forest management
in Poland are perceived as sustainable, the forest activities
may only be classied as the potential threat for the BF.
Recommendations for the majority of the BF habitats (i.e.
riparian, oak-hornbeam, coniferous and boggy forests), which
have not yet been transformed as a result of forest management,
clearly point out to the need for exclusion of such forests from
management activities. In the case of transformed stands (on
deciduous forest sites), slow stand restoration is recommended
towards adjusting species composition to site conditions.
5. Species diversity in the Białowieża Forest –
The BF is a ‘hot spot’ of Poland’s species diversity (Ja-
roszewicz 2010). Habitat mosaic and diversity, the presence
of old trees together with hollow and dying trees and dead-
wood abundance provide good life conditions for numero-
us species, including relict species characteristic for the old
primeval forests. Some of these have been preserved only in
the BF. It is impossible to completely describe biotic biodi-
versity of the BF; therefore, only selected groups of orga-
nisms are described in the following sections.
In the BF, macrofungal species diversity has been thus far
recognised at a range of levels. For the most part, the area of the
BNP has been studied in this respect. In the years 1987–1991,
the project CRYPTO was conducted in this region, and in one
forest division (no. 256), 913 species of macrofungi (Faliński,
Mułenko 1997) were found. Data on fungal species diversity
in the BF (mainly BNP) were gathered by many mycologists,
amongst others, Pilát (1950), Nespiak (1959), Orłoś (1960),
Skirgiełło (1960, 1998), Domański (1967), Bujakiewicz (1994)
and Karasiński et al. (2009). The results of mycological studies
carried out in the Forest were in some measure reviewed by
Karasiński et al. (2010) during the works on the nature mana-
gement plan for the BNP. Outside the Park’s borders, research
on fungi was carried out only in selected areas, and particularly,
in the nature reserves (Bujakiewicz 2002, 2003; Bujakiewicz,
Kujawa 2010). In the BF as a whole, most detailed studies
concerned polyporoid fungi (Niemelä 2013; Karasiński, Woł-
kowycki 2015). Up to date, 1,850 species of macrofungi have
been described in the BF (Kujawa, unpublished), that is, 43%
of fungal species reported from Poland. More than half (933) of
the fungal species in the BF are rare (included in Poland’s Red
List or found in one to three sites in the country or just lately
described – thus, not included in the lists of critical species)
or under legal protection. The group of rare species comprises
almost 200 species of fungi that have never been found on other
sites in the country (Kujawa, unpublished). The Forest’s my-
cobiota has not yet been fully recognised and potentially will
be broader, which is conrmed by the fact that new species of
macrofungi are reported every year during the cyclic exhibition
of fungi, held in the Białowieża village (e.g. Szczepkowski et
al. 2008, 2011; Gierczyk et al. 2013, 2014, 2015).
307A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
With regard to the species richness of polyporoid fungi, the
BF stands out from the rest of European forests. From the group
of 394 species of polyporales known in Europe, as many as 210
have been observed in the BF. This is almost 90% of all poly-
poroid fungi known in Poland (Karasiński, Wołkowycki 2015).
The above results demonstrate the uniqueness of the BF. At
the same time, the aggregation of many habitats of fungi, espe-
cially those associated with declining trees and deadwood, in
the areas under strict protection (mainly in the BNP) proves
that the passive protection of forest ecosystems is efcient for
safeguarding high diversity of fungal species associated with
forests. This especially concerns hemerophobic species, sen-
sitive to ecosystem changes because of management activities
carried out by man. There is a need for further research on the
status of species diversity of macrofungi in the BF.
5.2. Lichenised fungi (lichens)
The biota of lichenised fungi that occurs in the BF has
been quite well recognised. Information has been provided
by numerous studies carried out from the 1800s. A review
of earlier works (e.g. Błoński 1888; Krawiec 1938; Lece-
wicz 1954; Rydzak 1961) and wide-ranging information on
the distribution and ecological preferences of 309 species
of lichens occurring in the BF are available in the paper by
Cieśliński and Tobolewski (1988). The study area V-100 of
the Project CRYPTO conducted in the BNP (Protected Natu-
re Reserve, forest unit no. 256) was best described in terms
of the biota of lichenised fungi. Within the area of one forest
unit (140 ha), 164 species of lichens were identied (Cie-
śliński et al. 1995; Cieśliński, Czyżewska 1997).
In the register of lichenised fungi occurring in the BF
(Cieśliński 2010 and references), 450 lichen species are li-
sted (including 268 observed in BNP).
The updated list of about 500 species of lichenised fungi
and about 50 species of fungi living on lichens known from
the BF also comprises the information published in the recent
years (e.g. Matwiejuk, Bohdan 2011; Kukwa et al. 2012a,b;
Łubek, Jaroszewicz 2012; Guzow-Krzemińska et al. 2016;
Łubek, Kukwa 2016), as well as data on lichens obtained in
the project KlimaVeg (http://www.klimaveg.eu/), conducted
in V-100 study area (Łubek, Kukwa unpublished). Lichen
Table 1. List of natural habitats which are the objects of protection according to the Standard Data Form for the Natura 2000 the Białowieża
code Habitat name State of
Habitat area [ha] / share in the
total Nature 2000 area [%]
1. 3150 Natural eutrophic lakes with Magnopotamion or Hydrocharition - type
vegetation C12.63 / 0.02
2. 6230 Species-rich Nardus grasslands, on silicious substrates in mountain areas
(and submountain areas in Continental Europe) B 132.61 / 0.21
3. 6510 Lowland hay meadows (Alopecurus pratensis, Sanguisorba ofcinalis) B 524.13 / 0.83
4. 7140 Transition mires and quaking bogs B 18.94 / 0.03
5. 7230 Alkaline fens C 157.87 / 0.25
In total: 849.18 / 1.34
6. 9170 Galio-Carpinetum and Tilio-Carpinetum oak-hornbeam forests A 39 814.56 / 63.05
7. 91D0 Bog woodland B 2 746.92 / 4.35
8. 91E0 Alluvial forests with Alnus glutinosa and Fraxinus excelsior (Alno-Padi-
on, Alnion incanae, Salicion albae)A 12.63 / 0.02
Riparian mixed forests of Quercus robur, Ulmus laevis and Ulmus minor,
Fraxinus excelsior or Fraxinus angustifolia, along the great rivers (Ulme-
63.15 / 0.10
10. 91I0 Euro-Siberian steppic woods with Quercus spp. C 6.31 / 0.01
In total: 42 643.57 / 67.53
Natural habitats in total: 43 492.75 / 68.87
Explanations: A – excellent, B – good, C – medium or degraded
308 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
species that have been so far identied in the BF represent
about 30% of the biota of lichenised fungi reported from Po-
land and about 20% of the biota of non-lichenised fungi living
on lichens (e.g. Fałtynowicz 2003; Czyżewska, Kukwa 2009).
Back in the 1930s, the biota of lichenised fungi occurring
in the BF was characterised by the great abundance of foliose
macrolichens, such as Lobaria pulmonaria, that formed giant
thalli with fruiting bodies, overgrowing tree trunks and bran-
ches, including those of spruces. Also, thalli of fruticose and -
lamentous species of the genus Bryoria and Usnea and those of
the species such as Ramalina thrausta and Evernia divaricata
hung down from tree branches, creating a ‘primeval landscape’
(Krawiec 1938). Usnea longissima specimens with 1-m long
thalli were observed in the BNP still in the 1950s (Lecewicz
1954). Unfortunately, the end of the 20th century brought a
very strong change in the lichen biota across Poland and many
other European countries, as a result of increased air pollution
(e.g. Hawksworth et al. 1973; Kiszka 1977; Czyżewska 2003).
Lichens of the BF also suffered, despite of the considerable
distance from large industrial centres and main roads. Owing
to the impact of the long-range pollutions and contamination
derived from the small local sources (Malzahn 2009; Malza-
hn et al. 2009 and cited references), the species most sensitive
to acid rain and gaseous SO2 disappeared here (many species
from the genera Bryoria and Usnea) or their occurrence got
strongly reduced, for example, previously luxuriant Lobaria
pulmonaria was described as a very rare species, forming small
thalli without fruiting bodies (Cieśliński, Tobolewski 1988; see
also Gauslaa 1995 and cited references). In spite of these losses,
lichen biota of the BF is extremely rich in very rare species,
including those endangered in Poland (Cieśliński et al. 2006)
and protected by law (Rozporządzenie 2014). Thanks to the
presence of strictly protected forest areas in the Park, as well as
older forests in the nature reserves, this biota has a very large
share of red-listed lichens (52%), with approximately 40% of
the species in the highest threat categories (CR, EN, VU) (Cie-
śliński 2010). The BF is the most important refuge in Poland for
majority of these species. Amongst them, especially important
are the epiphytic and epixylic lichens, which are regarded as
primeval relicts (Cieśliński et al. 1996).
Relicts (species, indicators) of primeval forests are the rem-
nants of larger groups of species, typical components of pre-
viously widespread forest ecosystems, characterised by the
continuity of natural ecological processes, in which a key role
was played by generational turnover of tree species, shaping the
internal structure of forest community and habitats available for
lichens (Faliński 1986; Peterken 1996; Cieśliński et al. 1996).
The occurrence of primeval forest lichens depends on the pre-
sence of specic microniches, which are old and very old trees
of various species and wood in varying degrees of decompo-
sition and forms (dead standing or broken trunks without bark,
lower stumps, fallen logs, exposed root systems of fallen trees
Table 2. Selected conservation activities resulting from the plan of protection tasks for forest habitats and identied threats
Habitat code Protection tasks Potential threats*
9170 exclusion from the economic use all forest stands with the species in the composition of
10% at the age of 100 years or more on the habitat of oak-hornbeam subcontinental forest;
adjusting the composition of the tree stands to the composition in accordance with the
natural habitat; in the tree stands with domination of aspen, birch, pine and less fre-
quently spruce – in stands less than 100 year old
renaturisation of tree stands
elimination of invasive species
B02.04 – removal of dead and
B02 – forestry and tree plan-
tation, use of forests and
preventing the degradation of habitats by exclusion from economic activities patches
of habitat 91D0
reducing of maintenance and renewal of drainage ditches, except as necessary for the
maintenance of road and railway infrastructure
91E0 exclusion from the economic use all forest stands on the habitat 91E0
monitoring and removing of invasive species
B02.04 – removal of dead and
91F0 maintenance of proper water conditions in the catchment areas of forest rivers using
91I0 cuttings, limiting shading of the forest oor
* According to the “Reference list of pressures, threats and activities” included in the annex 5 of the Instruction for the Natura 2000 Standard Data Form,
version 2012.1, prepared by the General Directorate for Environmental Protection, available at http://natura2000.gdos.gov.pl/strona/nowy-element-3
309A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
and lying branches). The continuity of microclimate conditions
(especially high humidity) is very important for these species
(Cieśliński et al. 1996).
The distinguishing feature of the biota in the BF is a very
large group of such primeval forest micro- and macrolichens,
as well as local high frequency of many of them. Still quite
common are species such as Arthonia byssacea, Calicium ad-
spersum, Loxospora elatina, Opegrapha vermicellifera and
Pertusaria avida that belong to the category of endangered
species in the country (EN). It should be noted, however, that
their localities are concentrated mainly in the BNP and other
protected areas with the highest naturalness (Cieśliński, Czy-
żewska 2002; Cieśliński 2003; Czyżewska, Cieśliński 2003a,
b; Cieśliński 2009).
Data collected in the recent years (Golubkov et al. 2011;
Popławska 2012; Zalewska, Bohdan 2012, Bohdan 2014; Ma-
twiejuk, Bohdan 2014; Bohdan – unpublished; Zalewska et al.
– unpublished) indicate that populations of some macrolichens
critically endangered in Poland (Cieśliński et al. 2006) are
slowly regenerating in the BF, but almost exclusively within
the areas of the Park and nature reserves. A fairly large number
of the localities of Bryoria capillaris as well as fewer localities
of lichen species such as Ramalina thrausta, Usnea ceratina
and Lobaria amplissima (previously reported as L. virens – see
Kukwa et al. 2008) were observed. For the latter three species,
the BF is the last refuge, not only in Poland but also in the low-
lands of Central Europe. Furthermore, several other lichens
from the threat category CR, for example, Evernia divaricata,
Usnea glabrescens and U. orida (recently often included to
the species U. suboridana – Articus et al. 2002; Kukwa 2005;
Kościelniak 2007) have the greatest populations in the BF at the
country level. Some other species endangered in Poland are lo-
cally quite common, for example, Menegazzia terebrata (CR),
Thelotrema lepadinum (EN) and representatives of the genus
Cetrelia (EN), as well as Lobaria pulmonaria (EN) – someti-
mes forming large thalli (Ryś 2007; Paluch 2009).
Nearly all of the taxa described above have been placed on
the list covering a total of 71 species that are regarded as indi-
cators of primeval forests in the Polish Lowland (Czyżewska,
Cieśliński 2003c). The occurrence of species from this list is
applied in Poland in assessing the natural values of forests, as
in the case of indicator species used in other countries (e.g.
Rose 1976, Arup 1997, Coppins, Coppins, 2002; Rose, Cop-
pins, 2002; see also Kubiak 2013a,b, with references).
The main threat to rare, specialised forest lichens are chan-
ges in the structure of forest communities and the disconti-
nuity of microclimatic conditions related to timber acquisition
and regeneration of tree stands, regardless of the method used
(e.g. Rose 1992; Czyżewska 2003; Pykälä 2004; Scheidegger,
Werth 2009; Nascimbene et al. 2013 and cited references).
These rare lichen species can be best preserved when the
continuity of ecological processes is protected, that is, main-
taining the natural turnover of tree generations, which ensures
spontaneous uctuation of available substrates, along with the
natural dynamics of small, short-term gaps, which bring no
dramatic changes in microclimate conditions inside the forest.
In case of natural disturbances, such as windthrow or insect
outbreak, open spaces of different sizes are formed. In such
areas, the processes of slow, natural regeneration of forest
communities are accompanied by equally slow regeneration
of the biota of the specialised forest lichens. The condition for
the effectiveness of this process is the continuity of undistur-
bed areas around the sites with damaged communities. Thalli
of rare forest lichens inhabiting undisturbed areas serve as a
source of propagules for recolonisation of regenerated, ma-
ture phytocoenoses (Scheidegger, Werth 2009). The additio-
nal source, mainly for epixylic species, may be the thalli of
lichens growing on snags (dead broken trees), which usually
occur in the damaged and regenerating forest communities
(Czarnota 2012; Czarnota et al., unpublished).
Only large-area passive protection through a very long
period of time can assure regeneration of the natural lichen
biota in disturbed forest areas. The crucial role of the passive
protection in set-aside areas for the maintenance of rare fo-
rest lichens is emphasised in many studies (Cieśliński 2008;
Kościelniak 2008; Zalewska 2012; Kubiak 2013b, Nascim-
bene et al. 2013 – with cited references).
5.3. Vascular ora
The rst oristic notes on the vascular plant species of the
BF appeared in the monograph by Gilbert, dated 1791. The
19th century brought another development, amongst others,
by Brincken (1828), Gorski et al. (1829), Eichwald (1830)
and, nally, Paczoski (works carried out in 1897–1900)
as well as Błoński, Drymmer and Ejsmond (1888–1889).
During the World War I and after, much data about ora of
this region were provided by German and Polish botanists,
and particularly by J. Paczoski, the author of a masterpiece
publication ‘Forests of Białowieża’ dated from 1930 (So-
kołowski 1995). The ora of vascular plants occurring in
the Polish part of the Forest was most fully recognised by
Sokołowski (1995), conducting his research between 1961
and 1993, who recorded the presence of 1,017 species, re-
presenting 93 families and 428 genera. This number is about
the half of the vascular plant species occurring in the Polish
lowlands. From 1,017 species recorded by Sokołowski, 664
are the components of natural plant communities occurring
in the Forest, whereas the remaining 353 penetrated here as
a result of human activity (synanthropic species), counting
those ecologically (apothytes) or geographically alien.
The compactness of the BF complex and its high degree
of naturalness are reected in the number of the ancient wo-
odland indicator species. From 664 plant species determined
310 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
by Sokołowski (1995) as associated with natural plant com-
munities, 115 are those that show the afnity to ancient fo-
rests. These represent 17% of the vascular plant species that
are present in natural plant communities in the BF and 74%
of all the species of this category (as described by Dzwonko
and Loster 2001) that occur in Poland. This proportion is
considerably high, given that many of the plant species inc-
luded in the Polish list of ancient woodland indicator species
are plants occurring in mountainous regions, as well as those
whose geographical range does not cover the area of the BF.
Plants of this group can, therefore, be a good indicator of the
naturalness and continuity of centuries-old forest habitats
(sensu Peterken 1974) in large parts of the BF. Their afni-
ty to forests is a result of long-lasting evolution/adaptation
to habitats with low level of disturbance, operating in small
scale and with low frequency. Living under such conditions
caused that typical forest species did not evolve life history
traits that would allow them to for a quick escape, either in
space or in time (Hermy et al., 1999; Dzwonko, Loster 2001;
Whigham 2004). Hence, many of the so-called forest specia-
lists did not develop the ability to disperse over long distan-
ces, because about one-third of plant species of the herb
layer are autochores, myrmecochores (with seeds dispersed
by ants) or barochores (spread because of gravity) (Hermy
et al. 1999; Dzwonko, Loster 2001). Most herb layer species
produce short-lived seeds (longevity of less than 1–5 years)
and, in contrast to non-forest plants, do not form permanent
seed banks (Thompson et al. 1997). An expression of plant
adaptation to forest conditions is their longevity (the average
lifespan of forest perennials is 64 years – Ehrlén, Lehtilä
2002) and ability to clonal growth – typical for more than
80% of herb layer species (Klimeš et al. 1997). These fe-
atures are of great importance in the case of large-scale di-
sturbances with high frequency and intensity, because after
some time, these may lead to permanent loss of many forest
groundcover species. The adverse effects of disturbances
caused by human intervention in forest ecosystems, main-
ly due to the activities related to forest management, may
occur after several decades after the disturbance. At this
stage, stopping the progress of forest plant species loss may
be impossible (Tilman et al. 1994; Vellend et al. 2006).
Flora of the BF embraces many rare species, including those
of relict northern origin. The interim, subboreal nature of the
ora of this area is expressed by the presence of plant species
with a circumboreal distributional range, mostly representing
chorological Euro-Siberian and Central European elements,
although the representatives of sub-Atlantic and sub-Pontian
ora have also been observed here. Furthermore, several plant
species reach their eastern, south-western, western or north-we-
stern limits of their ranges in this region (Adamowski 2009).
According to Sokołowski (1995), about 35% of vascular
ora of the BF represents the previously mentioned synan-
thropic species. Some of them appeared here spontaneously,
others were introduced intentionally. Humans signicantly
alter habitats through their activities and create conditions for
the encroachment of plant species characteristic for places
completely transformed, such as clear-felled areas, mid-forest
roads, elds, meadows and glades. Amongst them, there are
native species (apophytes) and also geographically alien spe-
cies, including those from other regions of the country (e.g.
Carex brizoides and Acer pseudoplatanus) or dragged/intro-
duced from outside of Europe (e.g. Acer negundo, Quercus
rubra, Impatiens parviora) (Sokołowski 1995; Adamowski
2009). This certainly increases the overall species diversity of
vascular plants in the BF. Nevertheless, it also poses a huge
threat to native ora, as some alien species displace native
ones, changing the original species composition of plant com-
munities of this forest complex (Faliński 1998).
Up to date, nearly 10,000 species of insects have been
observed in the BF (Gutowski, Jaroszewicz 2001, 2004;
Gutowski et al. 2009; Gutowski unpublished) from 26,000
of species known from the Polish territory (Chudzicka, Ski-
bińska 2003; Razowski 1990, 1991a, 1991b, 1997a, 1997b).
Most probably, many more insect species occur here, as
every year brings new discoveries.
The orders such as true bugs, Hemiptera (652 species, re-
presenting 29% of Poland’s entomofauna); beetles, Coleopte-
ra (3,199 species, 51%); wasps, Hymenoptera (2,005 species,
33%); butteries and moths, Lepidoptera (1,609 species, 51%);
ies, Diptera (1,772 species, 26%) are outstanding in terms of
species richness. Given that the best identication concerns the
orders Coleoptera and Lepidoptera – as a result of more stu-
dies carried out on these groups both at the country and the BF
levels – as well as the fact that these orders are represented in
the Forest by more than half of the species known in Poland,
we can assume similar proportions with regard to the majori-
ty of other insect orders. The best-known Coleoptera families
include Apionidae (65% of Poland’s fauna in this group), jewel
beetles (Buprestidae, 55%), longhorn beetles (Cerambycidae,
65%), ladybirds (Coccinellidae, 68%), weevils (Curculionidae,
53%), false click beetles (Eucnemidae, more than 70%) and sap
beetles (Nitidulidae, 65%). These data were compiled by Gu-
towski and Jaroszewicz (2001, 2004), Gutowski et al. (2009)
and Jędryczkowski and Gutowski (2014). The results obtained
by these authors as well as summarised in their studies as result
of years of research carried out by numerous authors place the
BF at the forefront of not only Polish but also European refuges
of biodiversity of forest insects.
Saproxylic insects represent a group most typical for fore-
sts, which is exceptionally species rich and most vulnerable to
311A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
forest management (Grove 2002). A substantial part of their
life is directly associated with dying or dead trees (in various
forms and decomposition stages) or with diverse fungi and
insects colonising deadwood (Speight 1989). Apart from out-
standing insect species richness, when compared to other na-
tural forests, and especially those located within the European
Lowlands, the entomofauna of the BF is characterised by the
distinctively high representation of species considered relicts
of natural forest habitats, often referred to as relicts of pri-
meval forests. This term denotes the species that have disap-
peared from the majority of European forests, as a result of
their anthropogenic transformations because of considerable
depletion of forest ecosystems, attributable to the planned
forest management (in a relatively short period of time, i.e.
200–250 years of development of notional forest manage-
ment principles, and implemented with varying intensity in
different places in Europe, including Poland). The impove-
rishment of natural forests mainly refers to
• natural age structure of forest stands (currently, in most
managed forests, a signicant shortage, and sometimes the
lack of old trees, i.e. those achieving the age that individual
species can potentially reach, is observed – the assumed cut-
ting age of a given tree species, accepted in forestry practice,
has effectively eliminated the possibility of the formation of
forest natural age structure);
• natural spatial structure of forest stands, with the mo-
saic of forests affected by all kinds of disturbances, and the
areas covered with abundant herbaceous vegetation bloc-
king fast regeneration of tree stands or surfaces ‘held’ by
large herbivores ‘at the no-stand stage’ or the areas covered
by highly dispersed trees (currently, in the majority of ma-
naged forests, such spaces do not occur or occur for a short
time – the existing forest management principle for the full
use of habitat productive potential through rapid regenera-
tion has actually eliminated the possibility of the formation
of natural spatial structure of forest stands);
• natural species structure of forest stands, dynamically
changing at different rates, depending on the local factors
altering habitats that allow for the renewal and development
of various tree species – often undesirable from the econo-
mic point of view – including hornbeam, aspen and lime
(currently, in most managed forests, the accepted silvicultu-
ral and forest utilization principles rule out the possibility of
spontaneous formation of such a structure);
• abundance of deadwood, which is naturally present in
the forest – depending on its specic character and stand
uctuation stage – in a full range of forms (standing, lying,
suspended), as well as on the phase of wood decomposition
in a complete range of its dimensions, and the amount of
sunlight available – which all also refers to very old trees
(currently, in the majority of managed forests, sanitary,
harvest or tending cuttings, by denition have elimina-
ted the possibility of a steady stream of deadwood; dying
or fallen trees are removed from forests, which effectively
reduces or completely eliminates the creation of saproxylic
The presence of unique entomofauna in forests is directly
associated with the occurrence of the above-described ele-
ments of the ecological structure with its full diversity – typi-
cal for natural forest ecosystems. Amongst forest saproxylic
insects that endured in the BF’s numerous sites with pre-
served elements of natural ecological structure, as relicts of
primeval forests or the species associated with development
of natural forest microhabitats (e.g. old trees with the of hol-
lows shaped by lasting many decades processes of humica-
tion), the following ones deserve a special attention: Boros
schneideri (Boridae), Buprestis splendens (Buprestidae),
Leptura thoracica, Stictoleptura variicornis (Cerambyci-
dae), Ampedus melanurus, Lacon lepidopterus (Elateridae),
Otho sphondyloides (Eucnemidae), Lopheros lineatus (Ly-
cidae), Phryganophilus rucollis (Melandryidae), Pytho
kolwensis (Pythidae), Rhysodes sulcatus (Rhysodidae), Bius
thoracicus (Tenebrionidae) and Mycetoma suturale (Tetra-
tomidae). These Coleoptera species represent the ecological
group of saproxylic beetles that comprises more than 1,000
species observed in the BF. This demonstrates that the Fo-
rest is the main refuge of relict forest fauna in the North
European Plain (Gutowski, Jaroszewicz 2004; Gutowski
et al. 2009). Amongst saproxylic beetles observed here, as
many as 12 species included in Annexes II and IV of the EU
Habitats Directive occur here. Several species from other
ecological and systematic groups are also enlisted in the
Directive’s Annexes, for example, insects inhabiting aquatic
habitats and terrestrial open areas (e.g. dragonies, butter-
ies). Two hundred beetle species occurring in the BF are
included in the ‘Red List of threatened animals in Poland’
(Pawłowski et al. 2002).
There has been observed the presence of many species of
insects for whom the BF is one of a few or even the only site
in Central Europe. These are, amongst others, taiga species of
beetles (boreal and Siberian): Carphoborus cholodkovskyi,
Polygraphus punctifrons, Pityogenes saalasi, Orthotomicus
starki, Cryphalus saltuarius, Pityophthorus morosovi (Cur-
culionidae: Scolytinae); Acmaeops angusticollis, Evodinus
borealis, Stictoleptura variicornis, Leptura thoracica, Xylo-
trechus ibex, Mesosa myops and Monochamus sartor urus-
sovii (Cerambycidae). In the BF, insect species from other
zoogeographic regions have been observed, for example,
Aulonothroscus laticollis (Throscidae), Buprestis splendens,
Eurythyrea quercus, Agrilus pseudocyaneus (Buprestidae),
Nematodes lum (Eucnemidae), Pseudanostirus globicollis
(Elateridae), Alosterna ingrica (Cerambycidae) and Pachy-
tychius sparsutus (Curculionidae) (Gutowski, Jaroszewicz
2004). The Forest represents a unique refuge for rare Le-
312 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
pidoptera species, especially boreal ones – associated with
bogs, for example, Colias palaeno, Vacciiniina opilete, Bo-
loria eunomia, Euphydryas maturna and E. aurinia.
The occurrence of unique primeval entomofauna in the
BF is conditional on the continuous (from prehistoric times)
presence of standing dead trees and lying deadwood, as
well as the occurrence of natural stand uctuation stages,
including those of decay and regeneration. The latter is re-
lated to slowly progressing secondary succession hindered
by intense coverage of herbaceous vegetation on the areas
affected by natural disturbances (with weaker or stronger
effects) (e.g. Buchholz, Burakowski 1992). This is a result
of limited management of considerable portions of forest
ecosystems, including those not yet taken under protection
within the nature conservation system. Entomofauna species
richness, as well as the occurrence of species closely related
to the effects of spontaneously running ecological processes,
clearly indicates positive effects (for forest biodiversity) of
the exclusion or substantial reduction of forest management
based on the principles adopted in modern forestry.
Despite relatively good knowledge on the Forest’s ento-
mofauna when compared to other European forests, species
new to this area are still being discovered, including those
previously unknown in Poland, such as Acmaeops angusti-
collis (Cerambycidae) (Gutowski 1988), Isorhipis marmot-
tani (Eucnemidae) (Buchholz, Burakowski 1989), Ampedus
melanurus and A. suecicus (Elateridae) (Buchholz, Ossow-
ska 1998), Mordellochroa milleri (Mordellidae) (Kubisz
2000), Euplectus tholini (Staphylinidae) (Jałoszyński et al.
2005), Nacerdes carniolica (Mordellidae) (Gutowski et al.
2012) and Sepedophilus wankowiczi (Staphylinidae) (Szu-
jecki 2014). The BF is locus typicus for more than a dozen
species of Coleoptera, Diptera, Hymenoptera, Lepidoptera
and Mallophaga – recognised as new for the science (Di-
sney, Durska 1998; Okołów 2015; Gutowski unpublished).
In addition to the most valuable group of saproxylic insects,
a comparatively species-rich group of hygrophilous insects –
associated with waters and peat bogs – also occurs in the BF, for
example, dragonies (Odonata) are represented by 60 species
(83% of Poland’s dragony fauna). The group of xerophilic in-
sects is of slightly lower importance and less species rich (with
some exceptions) (Wanat 1994, 1999; Gutowski et al. 2009).
In the past decades, 153 breeding bird species were recor-
ded in the BF (Pugacewicz 1997), and these represent 67% of
contemporary breeding avifauna in Poland (Chodkiewicz et
al. 2015). Breeding avifauna of the BF comprises 30 species
listed in Annex I of the EU Birds Directive (Rowiński 2010).
The stands of the BF are exceptional in terms of extre-
mely rich, unique assemblage of forest and forest edge birds.
Breeding bird communities recorded here resemble the
structure of avifauna inhabiting undisturbed tropical forests
(a large number of species, low density, high levels of nest
predation) (Walankiewicz, 2002; Tomiałojć, Wesołowski,
2005; Czeszczewik et al. 2015). The number of species ne-
sting on sample plots established in 1975 on the area of only
187.5 ha is more than 80 (although for individual seasons
and plots this number is indeed smaller) (Tomiałojć et al.
1984; Wesołowski et al. 2015). Almost all European species
of woodpeckers nest in the Forest. The condition of the BF
avifauna is primarily due to a very large share of stands close
to primeval or old stands – originating from natural regene-
ration in forest areas disturbed or cut down nearly 100 years
ago (with many dead trees, including spruces and pines).
More than 40-year-long research on the assemblage of
birds in the BF has proved that it is an inherent ‘window to
the ecological past’ of European forests (Wesołowski, Ful-
ler 2012). The ecology of many bird species under the BF
conditions distinctively differs from the ecology of the same
species in heavily fragmented managed forests (Tomiałojć,
Wesołowski 2005; Wesołowski 2007). Studies conducted
in the BNP have shown that certain ways of nesting, unk-
nown or regarded as exceptional in other parts of the coun-
try and Europe, are relatively common here, and thus, our
views on evolution of the selection of a breeding site must
have been modied. This regards, for example, blackbird
Turdus merula and song thrush T. philomelos, frequently
nesting on naturally fallen trees (Tomiałołojć et al. 1984),
as well as robin Erithacus rubecula and blackbird – nesting
in tree cavities (Walankiewicz unpublished; Rowiński oral
communication). Wesołowski and Fuller (2012) published a
long list of differences between the BF and the UK forests
with regard to habitat preferences/nesting site choices of the
same bird species. This issue is of great importance, as in
ornithological literature, the presented depiction of ecology
of many European forest bird species has been shaped by the
results of numerous studies conducted on bird populations in
British, Dutch or Swedish forests and wooded lands heavily
transformed by man (Wesołowski 2007, Wesołowski, Fuller
2012; Jędrzejewska, Jędrzejewski 1998). Therefore, the BF
and the research conducted here is now the most important
point of reference for studies on ecology of birds in forests
of temperate Europe, as well as those in North America.
The protection of spontaneous ecological processes on-
going in bird assemblage is the most important priority in
research on ecology and behaviour of birds in the BF. For-
ty-year-long studies on avifauna conducted here enabled, for
example, description of a substantial increase in the total bird
densities in 1985–2001, followed by a downward phase (To-
miałojć et al. 1984; Wesołowski et al. 2015). The results obta-
ined by Walankiewicz (2002, 2006) showed that in hornbeam
and oak seeding years, the numbers of rodents and predators
313A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
uctuated, which caused changes in bird population numbers.
The ecological processes described above can be studied
only when as large as possible areas of naturally disturbed
old stands (e.g. by spruce bark beetle and other insect out-
breaks) remain free from human interference (Wesołowski,
2005; Wesołowski, Rowiński 2006; Czeszczewik et al. 2015).
The major threat to the BF avifauna and its ‘primeval ecolo-
gy’ is the transformation of fragments of natural or primeval, as
well as the ones affected by bark beetles (disturbed) tree stands,
which have endured in managed parts of the Forest, into a
‘simplied’ production forest. Forest management involves the
removal of old and dead trees and establishment of fenced re-
forestation areas with one or two dominant tree species, which
leads to the development of managed forest plantations typical
for Europe. The removal of individual dead trees (e.g. spruces
and pines) also constitutes a threat. Cavities in dead standing
tree trunks are important nesting sites for birds, sometimes used
for decades (Walankiewicz et al. 2014).
The strictly protected area within the BNP (less than 50
km2) is too small to sustain biologically viable populations
of woodpeckers, such as white-backed woodpecker Dend-
rocopos leucotos and three-toed woodpecker Picoides tri-
dactylus (Wesołowski 2005). The negative impact of forest
management on tree stands and associated avifauna was
demonstrated by comparing the groups of birds inhabiting
the BNP, nature reserves and managed parts of the Forest
(Czeszczewik et al. 2015). Likewise, the indicators of the
white-backed woodpecker population’s status have been de-
creasing with the increasing intensity of forest management
activities performed in the Forest’s stands (Walankiewicz et
al. 2011). As emphasised in the current recommendations in
the nature management plan of the BNP (Regulation 2014b),
hollow, dead and dying trees, as well as those fallen as a re-
sult of natural reasons, play an imperative role in the life of
numerous protected species of birds.
In conclusion, the best way to safeguard the ecological
processes along with the whole, extremely valuable assem-
blage of breeding birds is the passive protection of the whole
Polish part of the BF.
A distinct gradient in mammal species numbers has been
observed in Poland – from the greatest abundance in the south
to the lowest in north-eastern parts of the country (Ciecha-
nowski, Bogdanowicz 2014). Despite the location on Po-
land’s northern outskirts, the BF is remarkable in terms of
species-rich mammal community (approximately 60 species)
(Stachura et al. 2004; Rachwald, Ruczyński 2015). In ad-
dition to numerous species of the temperate zone, mammals
characteristic for the boreal zone occur here (whilst at the
same time they are absent in the central and southern parts of
the country), such as snow hare Lepus timidus (Gryz, Krauze-
Gryz 2014) and Laxmann’s shrew Sorex caecutiens (Pucek
2001b) – both considered as the post-glacial relicts. The BF
is the south-western limit of L. timidus range (Zbyryt et al.
2014), and in the case of S. caecutiens, the Forest constitutes
an islet in the far south, beyond the continuous range of the
species (van der Kooij et al. 2015). Notwithstanding the long-
term protection and good maintenance of forest habitats, BF
mammal fauna has not been able to resist the negative effects
of human activities and has been depleted of several species,
including brown bear Ursus arctos and European mink (Mu-
stela lutreola) (Stachura et al. 2004), in the past centuries.
In the BF, the protection of European bison Bison bonasus
is one of the main conservation objectives, as the Forest is the
most important refuge for this species. This is mainly a con-
sequence of its history – the last exterminated wild individuals
used to live here, and European bison population was reintro-
duced here, rst – thanks to breeding, and then as free-living
herds (Krasińska, Krasiński 2004). Modern scientic research
has challenged earlier beliefs, according to which bisons were
perceived as the animals strongly associated with forests. The
evolution of the species, dental morphology, behaviour, food
intake and microhabitat preferences indicate that it is a rumi-
nant species, associated with grassy open areas, rich in plant
species (Kerley et al. 2012; Bocherens et al. 2015). This is re-
ected, amongst others, in the BNP nature management plan,
which emphasises preservation of non-forest ecosystems as
feeding sites for European bisons (Regulation 2014b).
Thanks to considerably high proportion of stands of na-
tural character, as well as the presence of numerous dead
and dying trees in various stages of decomposition, both
standing and lying, mammals in the BF have numerous
natural shelters that are not present or available in limited
numbers within managed forests. Such shelters are crucial
for protected arboreal rodents, such as red squirrel Sciurus
vulgaris, common dormouse Muscardinus avellanarius, fo-
rest dormouse Dryomys nitedula and fat dormouse Glis glis
(Ściński, Borowski 2006, 2008; Czeszczewik et al. 2008). In
Poland, the two latter species have the status of near-threate-
ned (Pucek 2001a; Pucek, Jurczyszyn 2001).
Hollows, spaces under the protruding bark and cracks for-
med in decaying trunks of standing dying trees are also key
shelters for many species of bats – strictly protected Poland.
Fifteen bat species have been recorded in the BF (Stachura
et al. 2004; Rachwald, Ruczyński 2015). Shelter preferences
of common noctule Nyctalus noctula and lesser noctule N.
leisleri – species at high risk of extinction in Poland (Vo-
loshin 2001) – have been studied in detail. Noctules inhabit
both the hollows carved out by woodpeckers as well as the
holes in tree trunks formed as a result of wood decompo-
sition processes (as a rule, in oak and ash trees; less frequen-
314 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
tly, in alders, maples, hornbeams, pines and lime trees). The
trees with bat hiding places have a signicantly greater dia-
meter at breast height (DBH) (on an average 84 cm) when
compared to those growing nearby – not inhabited (average
DBH of 40 cm). The inhabited trees are also characterised
by a much older age, usually more than 160 years. Noctu-
les prefer to hide in dying trees (80%), over those alive or
completely dead. Unlike birds, bats chose the hollows situ-
ated high, on an average 18 m above the ground level (Ru-
czyński, Ruczyńska 2000; Ruczyński, Bogdanowicz 2005,
2008). Similar preferences for hiding places – mostly slots in
protruding bark, located in high dying trees – were observed
in western barbastelle Barbastelle barbastellus – one of the
species protected under Natura 2000. This bat species usual-
ly chooses hiding in the stands where forest management is
limited or absent (Russo et al., 2004), and individuals spe-
cimens return year after year to their favourable patches of
forest and even to the same trees (Hillen et al. 2010). For this
reason, in the nature management plan for the BNP, one of
the conditions to ensure the favourable conservation status
of B. barbastellus is the preservation of forest areas with a
high proportion of old trees as well as the presence of old
trees with hollows (Zarządzenie 2014b).
Dead or dying trees (including lying deadwood) play a
crucial role also in the life of larger mammal species. Decay-
ed stumps of naturally fallen trees are often used by badgers
Meles meles as a temporary shelter (Kowalczyk et al. 2004).
In 95% cases, pine martens Martes martes use hiding places
situated on trees; the females prefer the hollows and cracks
in the trunks to hide their offspring (Zalewski 1997). Wolves
Canis lupus – the priority species according to the Habitats
Directive – give birth to their young not only in burrows but
also in windthrows (Schmidt et al. 2008).
In turn, lynx (Lynx lynx) – also subject to the protection
under Natura 2000 – hunt most often in forest patches with
numerous fallen and wind-thrown trees that provide them
cover whilst approaching the prey (Podgórski et al. 2008).
Therefore, to protect this wild cat threatened with extinction
in Poland (Wolsan, Okarma 2001), it is crucial to safeguard
forest diversity and assure the presence of a large number
of dead trees, in particular – lying trunks and windthrows
(Schmidt et al. 2007). Hence, in the BF as a whole, preserva-
tion of the mosaic diversication of forest habitats structure,
typical for natural forests (fallen or broken trees, woodland
glades, along with natural regeneration with deadwood at
a level above 10% of stand stock) should be promoted in
line with the current recommendations of the nature mana-
gement plan for the BNP (Rozporządzenie 2014b). Protec-
tive measures for large carnivores in the Forest cannot be
implemented solely within the BNP, as lynx home ranges
and territories of wolf family groups are much larger than
the area of the Park (Jedrzejewska, Jędrzejewski 1998).
6. Future of the Białowieża Forest – Conclusions
The protection of natural processes (either strict or active
conservative) should predominate in the BF, because it al-
lows for, amongst others, preservation of adequate quantity
and quality of deadwood and old trees, indispensable for the
survival and development of numerous organisms (including
species endangered in Europe) (amongst others: Cieśliński et
al. 1996; Gutowski, Buchholz 2000; Czyżewska, Cieśliński
2003c; Gutowski et al. 2004; Cieśliński 2009; Kujawa 2009;
Bohdan 2014; Karasiński, Wołkowycki 2016). It is also im-
portant that the area of diverse protected environments should
be sufciently large to comprise all uctuation stages of tree
stands, thus providing microhabitats suitable for the develop-
ment and survival of organisms with very different ecological
requirements. Within the large forest, where the protection of
spontaneous natural processes is the main concern, there are
always open or semi-open areas, appearing as a result of the
natural disturbances (e.g. death of old trees, insect outbreak,
damage to hurricane winds, and frost or ood). Animal spe-
cies (in particular many invertebrates and birds), plants and
fungi are associated with these areas that give them an oppor-
tunity to nd suitable niches. It is obvious that in such forests,
species populations do not reach large numbers, but species
richness is at least the same as that in forest areas exploited
economically, where there is always more open space availa-
ble. The main reason behind this is evolutionary adaptation of
organisms associated with mid-forest open areas to rapid de-
tection and colonisation of such habitats. Open habitats within
forests, which once covered almost the entire continent, often
appeared for a relatively short time, in different places, some-
times distant from each other.
The lack of a solid, comprehensive management plan to
protect this precious area, along with recommendations in
the relevant documents regarding the protection of the Fo-
rest (see Section 3), as well as currently existing threats to its
nature, resulting, amongst others, from the fact that in many
managed parts of the Forest, timber harvest, ‘the protection
with the use of the methods of ecological engineering’ and
‘reconstruction of deciduous oak-hornbeam forests’ are
planned, all these underline the urgent need for a coherent
approach based on the following principles/priorities:
• the protection (strict or active conservative) of natural
processes taking place in forest communities. This approach
should include forest stands growing in deciduous habitats
with incompatible tree species composition (in particular, ex-
cessive representation of spruce) as a result of former forest
management activities. Reconstruction of such tree stands
should be performed by forces of nature. The above recom-
mendations should not apply to heliophilous oak commu-
nities, the degenerated patches of which endured in the BF
on the area of few hectares. In order to preserve the oristic
315A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
composition of this anthropozoogenic habitat (resulting from
cattle grazing), continuation of cattle grazing is required in the
forest. Abandonment of grazing leads to decrease in species
richness, hornbeam regeneration and natural return of species
composition characteristic for oak-hornbeam forest;
• the active protection of some non-forest communities
and enclaves created by man, which are now habitats of
protected species (clearings, previous repositories of wood,
gravel pits, etc.), in line with the recommendations of the
protection action plan for Natura 2000 site;
• limiting the timber harvesting to small areas, in order
to provide wood for local communities (as suggested in the
legislation projects proposed in 2006);
• lessening management activities in the Forest (also in
the aforementioned stands with species composition incompa-
tible with the site); in accordance with the recommendations
of the plan of protection tasks for Natura 2000 site as well as
those in the renomination dossier submitted to UNESCO.
In the debate about the future of the BF, the proponents of
its economic use have raised arguments that much of this forest
complex is already covered by different forms of protection.
In addition to the BNP, there is a dense network of nature re-
serves, where forests with ongoing natural processes are pro-
tected. Hence, the advocates of this approach believe that the
existing network of protected areas (i.e. the BNP and the na-
ture reserves) is sufcient to protect all the natural values of
the Forest as a whole, and for that reason, the remaining part
can be used economically. Furthermore, they emphasise the
fact that the strict protection of no more than 20% of the For-
est is sufcient to study the natural processes occurring in the
Forest (Brzeziecki 2016). Assuming the need for sustainable
management of forests, some scientists, including Jan Marek
Matuszkiewicz, make an allowance for articial regeneration
of the sites with removed spruce stands with various species
of trees (Hilszczański 2016). However, a cursory analysis of
the maps of the distribution of protected areas in the BF shows
that the continuation of the economic use of the rest of the For-
est will lead over time to a signicant isolation of the currently
preserved most valuable areas. Such a fate has befallen many
Polish forest complexes as well as those in the world. Most of
them are nameless forests while others, like those famous situ-
ated in the state of Wisconsin in the United States (Curtis 1956;
cit. after Burgess and Sharpe 1981) are a key example of human
folly and greed. For decades, scientic literature has been full
of examples of research on the fragmentation of forest cover
and its consequences for various organisms as well as ecologi-
cal processes (e.g. Helliwell 1976, Burgess, Sharpe 1981, Bur-
gess 1988; Harris, Silva-Lopez, 1992; Honnay et al. 2005). The
popular web search service Web of Science gives 26,090 results
for ‘forest fragmentation’ (access date June 21, 2016). In addi-
tion to climate change and biological invasions, habitat frag-
mentation is one of the three main factors causing the loss of the
world’s biotic diversity (Jackson, Sax 2010). This is the effect
of shrinking the area of the habitats suitable for survival along
with increasing the isolation of populations of various species
living in forests. This leads to a reduction of genetic variation
and an increase of genetic differentiation between populations
from different patches of the same forest (because of the genetic
drift, increased inbreeding and decreased gene ow between
populations spatially isolated from each other) (Honnay et al.
2005). Forest habitat fragmentation may adversely affect abun-
dance and diversity of various groups of organisms present in
the isolated forest patches. It can also cause an increase in forest
perimeter to forest area ratio, in other words: an enhancement
of the so-called edge effects. The area affected by the latter will
grow at the expense of the internal (core) forest area, which is
as a rule free of the edge of effect, and has different microcli-
mate that determines the existence of many forest species. With
the increasing range of edge effects and changes in microcli-
mate in the forest, the risk of invasion of interior forest habitats
by alien species (which naturally either do not occur there or
occur only temporarily and less frequently) increases. Micro-
climate changes may in fact give invasive species additional
chances to compete with forest species (Honnay et al. 2005 and
The processes described above, caused by forest cover
fragmentation and followed by intensied disturbances,
adversely affect the reproductive success of many forest spe-
cies. Some of these processes are initiated after a short time,
whilst the consequences of others are revealed after many
years. For certain groups of organisms, including herb layer
plants, the effects taking place in their disturbed environ-
ments, including those related to forest fragmentation, be-
come apparent after many decades. Forest plants, owing to
their longevity and clonal growth, outwardly show a redu-
ced sensitivity to habitat fragmentation, but in fact their re-
sponse is delayed in time. Consequently, currently observed
distribution of forest plants is not in equilibrium with the
present level of habitat fragmentation and disturbances in
their environment (Eriksson, Ehrlen 2001). Some species
will become extinct in the future, once a new equilibrium is
established in response to the disturbance in their habitats.
This phenomenon was described by Tilman et al. (1994) as
the extinction debt. It was demonstrated in the case of herb
layer species (Kolk, Naaf 2015; Naaf, Kolk 2015), epiphy-
tic forest lichens (Berglund, Jonsson, 2005; Ellis, Coppins
2007), meadow plants (Lindborg, Eriksson 2004) and some
animal species (Hanski, Ovaskainen 2002). In the case of
forest herb layer species of the temperate zone, the estimated
duration of the extinction debt ‘payoff’, that is, successive
species loss after the occurrence of disturbances, is estima-
ted to be about 100–250 years (Vellend et al. 2006).
Exploitation of the currently managed forests of the BF
triggers many not fully understood processes, some of which
316 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
may be irreversible in their consequences for wildlife. Un-
deniably, the slow and gradual transformation of subsequent
parts of the BF will lead to the situation that it will share the
fate of other temperate climate forests of Europe and North
America. The continuation of silvicultural techniques used
in commercial forests is destructive for the nature of this
region. The undesirable changes that have occurred here as
a result of forest management can sometimes be reversible
once the management activities are abandoned. The future
of the BF primarily depends on the changes in the way we
think about it. The endurance of this unique forest for future
generations, in the condition the least altered by humans,
can only be assured if we stop perceiving just the econo-
mic dimension of it (as the value obtained from wood raw
material) and start appreciating the value of its ecosystem
services, and if we limit the Forest’s use to the minimum
satisfying no more than the needs of the local community.
Conict of interest
The authors declare no potential conicts.
Acknowledgements and source of funding
The authors would like to thank the anonymous Revie-
wer for insightful comments and positive evaluation of the
The research was funded from the authors’ own sources.
Adamowski W. 2009. Flora naczyniowa, w: Białowieski Park Na-
rodowy. Poznać – Zrozumieć – Zachować (red. C. Okołów, M.
Karaś, A. Bołbot), Białowieski Park Narodowy, Białowieża,
59–72. ISBN 978-83-87054-68-7.
Arup U. 1997. Skoglig kontinuitet, in: Skyddsvärda lavar i syd-
västra Sverige (edit. U. Arup, S. Ekman, I. Kärnefelt, J.-E.
Mattsson). SBF-förlaget, Lund, 92–95. ISBN 91-972863-1-1.
Articus K., Mattsson J.E., Tibell L., Grube M., Wedin M. 2002.
Ribosomal DNA and ß-tubulin data do not support the sepa-
ration of the lichens Usnea orida and U. suboridana as dis-
tinct species. Mycological Research 4: 412–418. DOI 10.1017/
Battisse M. 1982. The Biosphere Reserve: A Tool for Environment
Conservation and management. Environmental Conservation
Berglund H., Jonsson B.G. 2005. Verifying an extinction debt among
lichens and fungi in northern Swedish boreal forests. Conservation
Biology 19: 338–348. DOI 10.1111/j.1523-1739.2005.00550.x.
BirdLife International. 2016. Important Bird and Biodiversity Area
factsheet: Białowieża Forest, http://www.birdlife.org [11.07.2016].
Blicharska M., Angelstam P. 2010. Conservation at risk: conict anal-
ysis in the Białowieża Forest, a European biodiversity hotspot. In-
ternational Journal of Biodiversity Science, Ecosystem Services &
Management 6(1): 68–74. DOI 10.1080/21513732.2010.520028.
Blicharska M., Van Herzele A. 2015. What a forest? Whose forest?
Struggles over concepts and meanings in the debate about the
conservation of the Białowieża Forest in Poland. Forest Policy
and Economics 57: 22–30. DOI 10.1016/j.forpol.2015.04.003.
Blicharska M, Bobiec A., Bohdan A., Buchholz L., Chylarecki P.,
Engel J., Falkowski M., Gutowski J.M., Jaroszewicz B., Kepel
A., Kujawa A., Latałowa M., Mikusiński G., Mysłajek R.W.,
Nowak S., Orczewska A., Skubała P., Stepaniuk M., Walankie-
wicz W., Wesołowski T., Zub K. 2016. Spór o przyszłość Pusz-
czy Białowieskiej. Las Polski 11: 8–9.
Błoński F. 1888. Spis roślin skrytopłciowych zebranych w 1887 r.
w Puszczy Białowieskiej. Pamiętnik Fizjograczny 8: 75–119.
Bobiec A. 2012. Białowieża Primeval Forest as a remnant of cul-
turally modied ancient forest. European Journal of Forest
Research 131: 1269–1285. DOI 10.1007/s10342-012-0597-6.
Bobiec A., Bobiec M. 2012. Inuence of spruce decline in stands
of the Białowieża National Park on natural oak regeneration.
Sylwan 156(4): 243–251.
Boch S., Prati D., Müller J., Socher S., Baumbach H., Buscot F.,
Gockel S., Hemp A., Hessenmöller D., Kalko E.K.V., Linsen-
mair K.E., Pfeiffer S., Pommer U., Schöning I., Schulze E.-D.,
Seilwinder C., Weisser W.W., Wells K., Fischer M. 2013. High
plant species richness indicates management-related disturbanc-
es rather than the conservation status of forests. Basic and Ap-
plied Ecology 14: 496–505. DOI 10.1016/j.baae.2013.06.001.
Bocherens H., Hofman-Kamińska E., Drucker D.G., Schmölcke
U., Kowalczyk R. 2015. European bison as a refugee species?
Evidence from isotopic data on early Holocene bison and
other large herbivores in Northern Europe. PLoS ONE 10(2):
e0115090. DOI 10.1371/journal.pone.0115090.
Bohdan A. 2014. Znaczenie ochrony biernej dla zachowania po-
rostów – reliktów lasów pochodzenia pierwotnego w Puszczy
Białowieskiej. Przegląd Przyrodniczy 25(4) 151–161.
Brzeziecki B. 2016. Trzeba dostrzec dynamikę Puszczy. Las Polski
Brzeziecki B., Pommerenin A., Miścicki S., Drozdowski S., Ży-
bura H. 2016. A common lack of demographic equilibrium
among tree species in Białowieża National Park (NE Poland):
evidence from long-term plots. Journal of Vegetation Science
27: 460–469. DOI 10.1111/jvs.12369.
Buchholz L., Burakowski B. 1989. Isorhipis marmottani (Bonvouloir,
1781) (Coleoptera, Eucnemidae) – nowy dla fauny Polski przed-
stawiciel goleńczykowatych. Przegląd Zoologiczny 33(1): 89–95.
Buchholz L., Burakowski B. 1992. Werykacja danych o występo-
waniu i nowe stanowiska Pseudanostirus globicollis (Germ.)
(Coleoptera, Elateridae) w Polsce. Wiadomości Entomologicz-
ne 11(2): 121–122.
Buchholz L., Ossowska M. 1998. Nowe dane o występowaniu
czterech mało znanych gatunków z rodziny sprężykowatych
(Coleoptera: Elateridae), w niektórych rejonach Europy Środ-
kowej. Wiadomości Entomologiczne 17(1): 21–36.
Bujakiewicz A. 1994. Macrofungi in the alder forests of the
Białowieża National Park. Mycologia Helvetica 6(2): 57–76.
Bujakiewicz A 2002. New, rare and endangered fungi in the Białow-
ieża Primeval Forest. Polish Botanical Journal 47: 113–124.
317A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
Bujakiewicz A. 2003. Puszcza Białowieska ostoją rzadkich i za-
grożonych grzybów wielkoowocnikowych. Parki Narodowe i
Rezerwaty Przyrody 22(3): 323–346.
Bujakiewicz A., Kujawa A. 2010. Grzyby wielkoowocnikowe wy-
branych rezerwatów przyrody Puszczy Białowieskiej. Parki
Narodowe i Rezerwaty Przyrody 29(1): 3–26.
BULiGL 2015. Ekspertyza na potrzeby aneksu do Planu Urządze-
nia Lasu Nadleśnictwa Białowieża zawierająca ocenę stanu
lasu oraz określająca zakres niezbędnych działań gospodarczo
-ochronnych dla zachowania drzewostanów świerkowych. Biuro
Urządzania Lasu i Geodezji Leśnej Oddział w Białymstoku.
Burgess R.L. 1988. Community organization: effects of landscape
fragmentation. Canadian Journal of Botany 66: 2687–2690.
Burgess R.L., Sharpe D.M. (edit.) 1981. Forest island dynamics in
man-dominated landscapes. Springer-Verlag, New York, Hei-
delberg, Berlin, 320 s. ISBN 978-0-387-90584-6.
Catford J.A., Daehler C.C., Murphy H.T., Sheppard A.W., Hardes-
ty B.D., Westcott D.A., Rejmánek M., Bellingham P.J., Pergl J.,
Horvitz C.C., Hulme P.E. 2012. The intermediate disturbance
hypothesis and plant invasions: Implications for species richness
and management. Perspectives in Plant Ecology, Evolution and
Systematics 14: 231–241. DOI 10.1016/j.ppees.2011.12.002.
Chodkiewicz T., Kuczyński L., Sikora A., Chylarecki P., Neubauer
G., Ławicki Ł., Stawarczyk T. 2015. Ocena liczebności popu-
lacji ptaków lęgowych w Polsce w latach 2008–2012. Ornis
Polonica 56: 149–189.
Chudzicka E., Skibińska E. 2003. Różnorodność gatunkowa
– zwierzęta, w: Różnorodność biologiczna Polski (red. R.
Andrzejewski, A. Weigle), Narodowy Fundusz Ochrony Śro-
dowiska, Warszawa, 93–138. ISBN: 83-85908-75-7.
Ciechanowski M., Bogdanowicz W. 2014. Mammalia, w: Fauna Pol-
ski. Charakterystyka I wykaz gatunków. Tom IV (red. W. Bogda-
nowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska). Muzeum i Instytut
Zoologii PAN, Warszawa, 429–517. ISBN 9788388147142.
Cieśliński S. 2003. Atlas rozmieszczenia porostów (Lichenes) w Pol-
sce północno-wschodniej. Phytocoenosis 15 (N.S.), Supplemen-
tum Cartographie Geobotanicae 15: 1–430.
Cieśliński S. 2008. Znaczenie ochrony rezerwatowej dla zacho-
wania bioty porostów (Ascomycota lichenisati) w Puszczy
Kozienickiej. Studia i Materiały Centrum Edukacji Przyrodni-
czo-Leśnej w Rogowie 3(19): 99–109.
Cieśliński S. 2009. Porosty, w: Białowieski Park Narodowy. Po-
znać – Zrozumieć – Zachować. (red. C. Okołów, M. Karaś,
A. Bołbot), Białowieski Park Narodowy, Białowieża, 73–86.
Cieśliński S. 2010. Wykaz gatunków porostów (grzybów zlicheni-
zowanych) Puszczy Białowieskiej (NE Polska). Parki Narodo-
we i Rezerwaty Przyrody 29(2): 3–39.
Cieśliński S., Czyżewska K. 1997. Lichenes, in: Cryptogamous
plants in the forest communities of Białowieża National Park.
Ecological Atlas (Project CRYPTO 4), (edit. J.B. Faliński, W.
Mułenko), Phytocoenosis 9 (N.S.), Supplementum Cartograph-
iae Geobotanicae 7: 123–163.
Cieśliński S., Czyżewska K. 2002. Porosty Puszczy Białowieskiej
na tle innych kompleksów leśnych w Polsce północno-wschod-
niej. Kosmos 51(4): 443–451.
Cieśliński S., Czyżewska K., Fabiszewski J. 2006. Red list of the li-
chens in Poland, in: Red list of plants and fungi in Poland. (edit.
Z. Mirek, K. Zarzycki, W. Wojewoda, Z. Szeląg). W. Szafer In-
stitute of Botany Polish Academy of Science, Kraków, 71–89.
Cieśliński S., Czyżewska K., Faliński J.B., Klama H., Mułenko W.,
Żarnowiec J., 1996. Relicts of the primeval (virgin) forest. Rel-
ict phenomena, in: Cryptogamous plants in the forest commu-
nities of Białowieża National Park (Project CRYPTO 3). (edit.
J.B. Faliński, W. Mułenko). Phytocoenosis 8 (N.S.) Archiwum
Geobotanicum 6: 197–216.
Cieśliński S., Tobolewski Z. 1988. Porosty (Lichenes) Puszczy
Białowieskiej i jej zachodniego przedpola. Phytocoenosis 1 (N.
S.), Supplementum Cartographiae Geobotanicae 1: 1–216.
Connell J.H. 1978. Diversity in tropical rain forests and coral reefs.
Science 199(4335): 1302–1310.
Coppins A.M., Coppins B.J. 2002. Indices of ecological continuity
for woodland epiphytic lichen habitats in the British Isles. Brit-
ish Lichen Society, London, 37 s. ISBN: 0 9540418 44.
Czarnota P. 2012. Lichen protection needs natural forest distur-
bances – examples from some Polish Western Carpathians na-
tional parks, in: Lichen protection – protected lichen species.
(edit. L. Lipnicki), Sonar Literacki, Gorzów Wielkopolski:
53–66. ISBN: 978-83-63189-19-8.
Czarnota P., Zalewska A., Szymczyk R. 2013. Unpubl. Importance
of post-bark beetle Norway spruce snags for the protection of
lichen diversity in the Western Carpathians.
Czerepko J. 2008. A long-term study of successional dynamics
in the forest wetlands. Forest Ecology and Management 255:
630–642. DOI 10.1016/j.foreco.2007.09.039.
Czeszczewik D., Walankiewicz W., Stańska M. 2008. Small mam-
mals in nests of cavity-nesting birds: Why should ornitholo-
gists study rodents? Canadian Journal of Zoology 86: 286–293.
Czeszczewik D., Zub K., Stanski T., Sahel M., Kapusta A., Walan-
kiewicz W. 2015. Effects of forest management on bird assem-
blages in the Białowieża Forest, Poland. iForest 8: 377–385.
Czyżewska K. 2003. Ocena zagrożenia bioty porostów Pol-
ski. Monographiae Botanicae 91: 241–249. DOI 10.5586/
Czyżewska K., Cieśliński S. 2003a. Czerwona lista porostów za-
grożonych w Puszczy Białowieskiej. Monographiae Botanicae
91: 107–119. DOI 10.5586/mb.2003.006.
Czyżewska K., Cieśliński S. 2003b. Regionalne czerwone listy
porostów zagrożonych. Monographiae Botanicae 91: 51– 62.
Czyżewska K., Cieśliński S. 2003c. Porosty – wskaźniki niżowych
lasów puszczańskich. Monographiae Botanicae 91: 223–239.
Czyżewska K., Kukwa M. 2009. Lichenicolous fungi of Poland.
A catalogue and key to species, in: Biodiversity of Poland 11.
(edit. Z. Mirek). W. Szafer Institute of Botany, Polish Academy
of Sciences, Kraków, 1–133.
Denisiuk Z., Witkowski Z. 1990. Rezerwaty Biosfery w Polsce.
Białowieski Park Narodowy. Komitet Naukowy przy Prezy-
dium PAN „Człowiek i Środowisko”, Zakład Ochrony Przy-
318 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
rody i Zasobów Naturalnych PAN. Ossolineum, 54 s. ISBN
Disney R.H.L., Durska E. 1998. A new genus and new species of
Phoridae from Poland. European Journal of Entomology 95:
Domański S. 1967. Specyka mikoory nadrzewnej Białowieskie-
go Parku Narodowego ze szczególnym uwzględnieniem grzy-
bów z rzędu Aphyllophorales. Sylwan 111(1): 17–27.
Drozdowski S., Brzeziecki B., Żybura H., Żybura B., Gawron L.,
Buraczyk W., Zajączkowski J., Bolibok L., Szeligowski H.,
Bielak K., Widawska Z. 2012. Wieloletnia dynamika starod-
rzewów w zagospodarowanej części Puszczy Białowieskiej:
gatunki ekspansywne i ustępujące. Sylwan 156(9): 663−671.
Dzwonko Z., Loster S. 2001. Wskaźnikowe gatunki roślin starych
lasów i ich znaczenie dla ochrony przyrody i kartograi roślin-
ności. Prace Geograczne 178: 119–132.
Ehrlén J., Lehtilä K. 2002. How perennial are perennial plants?
Oikos 98: 308–322. DOI 10.1034/j.1600-0706.2002.980212.x.
Ellis C.J., Coppins B.J. 2007. 19th century woodland structure controls
stand-scale epiphyte diversity in present-day Scotland. Diversity and
Distribution 13: 84–91. DOI 10.1111/j.1366-9516.2006.00310.x.
Eriksson O., Ehrlén J. 2001. Landscape fragmentation and the vi-
ability of plant populations, in: Integrating Ecology and Evo-
lution in a Spatial Context. 14th Special Symposium of the
British Ecological Society. (edit. J. Silvertown, J. Antonovics)
Blackwell Science, Oxford, 423 s. ISBN 9780521549332.
Faliński J.B. 1986. Vegetation dynamics in temperate lowland pri-
meval forest. Ecological studies in Białowieża forest. Geobota-
ny 8: 1–357. Dr Junk Publishers, Dordrecht-Boston-Lancaster.
Faliński J.B. 1998. Invasive alien plants, vegetation dynamics
and neophytism. Phytocoenosis 10 (N.S.) Supplementum
Carthographiae Geobotniacae 9: 163–187.
Faliński J.B., Mułenko W. (red.). 1997. Cryptogamous plants in the
forest communities of Białowieża National Park. Ecological
Atlas (Project CRYPTO 4). Phytocenosis 9 (N.S.), Supplemen-
tum Cartographie Geobotanicae.
Fałtynowicz W. 2003. Lichenes, lichenicolous and allied fungi of
Poland. An annotated checklist, in: Biodiversity of Poland 1
(red. Z. Mirek). W. Szafer Institute of Botany, Polish Academy
of Sciences, Kraków, 1–435. ISBN 83-89648-06-7.
Gauslaa Y. 1995. The Lobarion, an epiphytic community of ancient
forests threatened by acid rain. Lichenologist 27: 59–76. DOI
Gierczyk B., Kujawa A., Szczepkowski A., Karasiński D. 2014.
Materiały do poznania mycobioty Puszczy Białowieskiej.
Przegląd Przyrodniczy 25(1): 3–36.
Gierczyk B., Kujawa A., Szczepkowski A. 2015. XX Jubileuszowa
wystawa grzybów Puszczy Białowieskiej. Materiały do pozna-
nia mykobioty Puszczy Białowieskiej. Przegląd Przyrodniczy
Gierczyk B., Szczepkowski A., Kujawa A. 2013. XVIII Wystawa
Grzybów Puszczy Białowieskiej. Parki Narodowe i Rezerwaty
Przyrody 32(2): 88–112.
Golubkov V., Bohdan A., Popławska M. 2011. Nowe, rzadkie i
interesujące gatunki porostów Białowieskiego Parku Narodo-
wego. Parki Narodowe i Rezerwaty Przyrody 30(3–4): 15–26.
Grove S.J. 2002. Saproxylic insect ecology and the sustainable man-
agement of forests. Annual Review of Ecology and Systematics
33: 1–23. DOI 10.1146/annurev.ecolsys.33.010802.150507.
Gryz J., Krauze-Gryz D. 2014. Nowe stwierdzenie zająca bielaka
Lepus timidus w Puszczy Białowieskiej. Chrońmy Przyrodę
Ojczystą 70(3): 283–284.
Gutowski J.M. 1988. Acmaeops angusticollis (Gebler, 1833) (Co-
leoptera, Cerambycidae) w Polsce. Polskie Pismo Entomolo-
giczne 58(2): 493–496.
Gutowski J.M., Bobiec A., Pawlaczyk P., Zub K. (red.). 2004. Dru-
gie życie drzewa. WWF Polska, Warszawa – Hajnówka, 245 s.
Gutowski J.M., Buchholz L. 2000. Owady leśne – zagrożenia i
propozycje ochrony. Wiadomości Entomologiczne 18, Supl. 2:
Gutowski J.M., Czachorowski S., Górski P., Wanat M. 2009. XI Bez-
kręgowce, w: Białowieski Park Narodowy, Poznać-Zrozumieć
-Zachować. (red. C. Okołów, M. Karaś, A. Bołbot). Białowieski
Park Narodowy, Białowieża, 161–176. ISBN 9788387054687.
Gutowski J.M., Jaroszewicz B. (red.). 2001. Katalog fauny Pusz-
czy Białowieskiej. Instytut Badawczy Leśnictwa, Warszawa,
403 s. ISBN 83-87647-22-5.
Gutowski J.M., Jaroszewicz B. 2004. Puszcza Białowieska jako
ostoja europejskiej fauny owadów. Wiadomości Entomologicz-
ne 23, Supl. 2: 67–87.
Gutowski J.M., Kubisz D., Sućko K. 2012. Nacerdes carniolica
(Gistel, 1834) (Coleoptera: Oedemeridae) – nowy chrząszcz dla
polskiej fauny. Wiadomości Entomologiczne 31(4): 267–273.
Guzow-Krzemińska B., Czarnota P., Łubek A., Kukwa M. 2016.
Micarea soralifera sp. nov., a new sorediate species in the M.
prasina group. Lichenologist 48(3): 161–169. DOI 10.1017/
Hanski I., Ovaskainen O. 2002. Extinction debt at extinc-
tion threshold. Conservation Biology 16(3): 666–673. DOI
Harris L.D., Lopes G.S. 1992. Forest fragmentation and the con-
servation of biological diversity. in: Conservation Biology. The
theory and practice of nature conservation, preservation and
management. (edit. P.L. Fiedler, K.J. Subodh). Chapmann and
Hall. New York and London, 196–237. ISBN 0 412 01961 2.
Hawksworth D.L., Rose F., Coppins B.J. 1973. Changes in the li-
chen ora of England and Wales attributable to pollution by
sulphur dioxide, in: Air pollution and lichens. (edit. B.W. Ferry,
M.S. Baddeley, D.L. Hawksworth) Athlone Press, London,
331–367. ISBN 0485111403.
Hedemann O. 1939. Dzieje Puszczy Białowieskiej w Polsce prze-
drozbiorowej (w okresie do 1798 roku). Instytut Badawczy
Lasów Państwowych, Rozprawy i Sprawozdania, Seria A, 41,
Helliwell D.R. 1976. The effects of size and isolation on the con-
servation value of wooded sites in Britain. Journal of Biogeog-
raphy 3: 213–221.
Hermy M., Honnay O., Firbank L., Grashof-Bokdam C., Lawesson
J. 1999. An ecological comparison between ancient and other
forest plant species of Europe, and the implications for forest
conservation. Biological Conservation 91: 9–22. DOI 10.1016/
319A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
Hillen J., Kiefer A., Veith M. 2010. Interannual delity to roosting hab-
itat and ight paths by female western barbastelle bats. Acta Chi-
ropterologica 12(1): 187–195. DOI 10.3161/150811010X504680.
Hilszczański J. 2016. Blokada etatu cięć to był strzał w stopę.
Rozm. przepr. B. Grabowska. Głos Lasu 2: 8–13.
Honnay O., Jacquemyn H., Bossuyt B., Hermy M. 2005. Forest
fragmentation effects on patch occupancy and population via-
bility of herbaceous plant species. New Phytologist 166: 723–
36. DOI 10.1111/j.1469-8137.2005.01352.x.
Jackson S.T., Sax D.F. 2010. Balancing biodiversity in a changing
environment: extinction debt, immigration credit and species
turnover. Trends in Ecology and Evolution 25(3): 153–160.
Jałoszyński P., Gawroński R., Kaźmierczak M., Gutowski J.M.
2005. Nowe dla Polski i rzadkie chrząszcze z rodzaju Euplec-
tus Leach (Coleoptera: Staphylinidae: Pselaphinae). Wiado-
mości Entomologiczne 24(3): 147–152.
Jaroszewicz B. 2010. Charakterystyka przyrodnicza i historia
Puszczy Białowieskiej i jej przedpola, w: Z Mazowsza na Po-
lesie i Wileńszczyznę. Zróżnicowanie i ochrona szaty roślinnej
pogranicza Europy Środkowej i Północno-Wschodniej (red.
A. Obidziński). Polskie Towarzystwo Botaniczne, Warszawa,
213–223. ISBN 83-86292-78-4.
Jaroszewicz B., Bobiec A., Eycott A.E. 2016. Lack of demographic
equilibrium indicates natural, large-scale forest dynamics, not a
problematic forest conservation policy – a reply to Brzeziecki et
al. Journal of Vegetation Science DOI 10.1111/vs.12458 (in press).
Jaskanis J. 2012. Wodzowskie kurhany kultury wielbarskiej na
Podlasiu. Muzeum Podlaskie w Białymstoku, Białystok. ISBN
Jędryczkowski W.B., Gutowski J.M. 2014. Biedronkowate (Co-
leoptera: Coccinellidae) Puszczy Białowieskiej. Wiadomości
Entomologiczne 33(3): 200–213.
Jędrzejewska B., Jędrzejewski W. 1998. Predation in vertebrate
communities. The Białowieża Primeval Forest as a case study.
Springer-Verlag, Berlin, 452 s. ISBN 978-3-642-08384-6.
Karasiński D., Kujawa A., Piątek M., Ronikier A., Wołkowycki
M. 2009. Contribution to biodiversity assessment of European
primeval forests: new records of rare fungi in the Białowieża
Forest. Polish Botanical Journal 54(1): 55–97.
Karasiński D., Kujawa A., Szczepkowski A., Wołkowycki M.
2010. Plan Ochrony Białowieskiego Parku Narodowego. Ope-
rat ochrony gatunków grzybów. Maszynopis.
Karasiński D., Wołkowycki M. 2015. An annotated and illustrated
catalogue of Polypores (Agaricomycetes) of the Białowieża
Forest (NE Poland). Polish Botanical Journal 60(2): 217–292.
Keczyński A. 2002. Wpływ kornika drukarza na drzewostany ob-
szaru ochrony ścisłej Białowieskiego Parku Narodowego. Ko-
smos 257: 471–474.
Keczyński A. 2005. Zmiany struktury drzewostanów wykształco-
nych na glebach hydrogenicznych – na przykładzie wybranych
powierzchni badawczych w Białowieskim Parku Narodowym.
Leśne Prace Badawcze 4: 87–102.
Keczyński A. 2007. Regeneracja grądu Tilio-Carpinetum Tracz.
1962 w następstwie dawnego użytkowania lasu w Białowie-
skim Parku Narodowym. Sylwan 1: 58–65.
Kerley G.I.H., Kowalczyk R., Cromsigt J.P.G.M. 2012. Conserva-
tion implications of the refugee species concept and the European
bison: king of the forest or refugee in a marginal habitat? Eco-
graphy 35: 519–529. DOI 10.1111/j.1600-0587.2011.07146.x.
Kiszka J. 1977. Wpływ emisji miejskich i przemysłowych na orę
porostów (Lichenes) Krakowa i Puszczy Niepołomnickiej.
Prace Monograczne Wyższej Szkoły Pedagogicznej w Krako-
wie 19: 1–133.
Klimeš L., Klimešova J., Hendriks R., van Groenendael J. 1997.
Clonal plant architecture: a comparative analysis of form and
function, in: The Ecology and Evolution of Clonal Plants. (edit.
H. de Kroon, J. van Groenendael). Backhuys Publishers, Leid-
en, The Netherlands, 453 s. ISBN 90-73348 -73-0.
Kolk J., Naaf T. 2015. Herb layer extinction debt in highly fragmented
temperate forests – Completely paid after 160 years? Biological
Conservation 182: 164–172. DOI 10.1016/j.biocon.2014.12.004.
Kościelniak R. 2007. Usnea orida – threatened species of rich
biotopes in the Polish Eastern Carpathians. Acta Mycologica
42(2): 281–286. DOI 10.5586/am.2007.031.
Kościelniak R. 2008. Znaczenie lasów o charakterze pierwotnym
i naturalnym dla zachowania różnorodności gatunkowej poro-
stów w Bieszczadach. Roczniki Bieszczadzkie 16: 67–76.
Kowalczyk R., Zalewski A., Jędrzejewska B. 2004. Seasonal and
spatial pattern of shelter use by badgers Meles meles in Białow-
ieża Primeval Forest (Poland). Acta Theriologica 49: 75–92.
Krasińska M., Krasiński Z.A. 2004. Żubr. Monograa przyrodni-
cza. SFP Hajstra, Warszawa-Białowieża.
Krawiec F. 1938. Materiały do ory porostów północno-wschod-
niej Polski. Sprawozdania Komisji Fizjogracznej Polskiej
Akademii Umiejętności 71: 65–82.
Krzyściak-Kosińska R., Arnolbik V., Antczak A. 2012 “Belovezhs-
kaya Pushcha / Białowieża Forest” world heritage site (33 bis)
proposed modication of the criteria and boundaries change of
the name of the property. Nomination Dossier to the UNESCO
for the Inscription on the World Heritage List. http://whc.unes-
Kubiak D. 2013a. Porosty jako wskaźniki ciągłości ekologicz-
nej zbiorowisk leśnych, w: Biologiczne metody oceny stanu
środowiska. Tom 1. Ekosystemy lądowe. (red. M. Dynow-
ska, H. Ciecierska). Wyd. Mantis, Olsztyn, 125–151. ISBN
Kubiak D. 2013b. The signicance of old-growth forests in main-
taining lichen diversity – an example from the remnants of the
Mazovian Forest. Forest Research Papers 74(3): 245–255.
Kubisz D. 2000. Mordellochroa milleri Emery (Mordellidae),
Anaspis bohemica Schilsky (Scraptiidae) i Corticeus bi-
coloroides (Roubal) (Tenebrionidae) – nowe dla fauny Polski
gatunki chrząszczy (Coleoptera: Tenebrionoidea). Wiadomości
Entomologiczne 19(1): 9–14.
Kuijper, D.P.J., Jędrzejewska, B., Brzeziecki, B., Churski, M., Ję-
drzejewski, W., Żybura, H. 2010. Fluctuating ungulate density
shapes tree recruitment in natural stands of the Białowieża Pri-
meval forest, Poland. Journal of Vegetation Science 21: 1082–
1098. DOI 10.1111/j.1654-1103.2010.01217.x.
Kujawa A. 2009. Grzyby wielkoowocnikowe, w: Białowieski Park
Narodowy. Poznać – zrozumieć – zachować. (red. C. Okołów,
320 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
M. Karaś, A. Bołbot). Białowieski Park narodowy, Białowieża,
87–110. ISBN 9788387054687.
Kukwa M. 2005. Nowe stanowiska rzadkich i interesujących poro-
stów na Pomorzu Gdańskim. Acta Botanica Cassubica 5: 95–111.
Kukwa M., Łubek A., Szymczyk R., Zalewska A. 2012a. Seven
lichen species new to Poland. Mycotaxon 120: 105–118. DOI
Kukwa M., Pietnoczko M., Czyżewska K. 2012b. The lichen family
Parmeliaceae in Poland II. The genus Cetrelia. Acta Societatis Bo-
tanicorum Poloniae 81(1): 43–52. DOI 10.5586/asbp.2012.007.
Kukwa M., Schiefelbein U., Czarnota P., Halda J., Kubiak D., Pal-
ice Z., Naczk A. 2008. Notes on some noteworthy lichens and
allied fungi found in the Białowieża Primeval Forest in Poland.
Bryonora 41: 1–11.
Kwiatkowski W. 1994. Krajobrazy roślinne Puszczy Białowieskiej.
Phytocoenosis 6. Supplementum Cartographiae Geobotanicae
Latałowa M., Zimny M., Jędrzejewska B., Samojlik T. 2015.
Białowieża Primeval Forest: a 2000-year interplay of environ-
mental and cultural forces in Europe’s best preserved temperate
woodland, in: Europe’s changing woods and forests: From wild-
wood to managed landscapes. Chapter 17. (edit. K.J. Kirby, C.
Watkins) CAB International, 243–264. ISBN 9781780643373.
Latałowa M., Zimny M., Pędziszewska A., Kupryjanowicz M.
2016. Postglacjalna historia Puszczy Białowieskiej – roślin-
ność, klimat i działalność człowieka. Parki Narodowe i Rezer-
waty Przyrody 35(1) (w druku).
Lecewicz W. 1954. Porosty Białowieży. Fragmenta Floristica et
Geobotanica 1(2): 38–47.
Lindborg R., Eriksson O. 2004. Historical landscape connectivity
affects present plant species diversity. Ecology 85: 1840–1845.
Łubek A., Jaroszewicz B. 2012. New, rare and noteworthy spe-
cies of lichens and lichenicolous fungi from Białowieża For-
est. Polish Journal of Natural Sciences 27: 275–287. ISBN
Łubek A., Kukwa M. 2016. Grzyby naporostowe w zbiorowiskach
leśnych Białowieskiego Parku Narodowego, w: Botanika –
tradycja i nowoczesność. Streszczenia referatów i plakatów
57. Zjazdu Polskiego Towarzystwa Botanicznego, Lublin 27
czerwca – 3 lipca 2016 (red. E. Szczuka, G. Szymczak, M.
Śmigała, R. Marciniec), 205–206. ISBN 978-83-86292-58-5.
Malzahn E. 2009. Biomonitoring środowiska leśnego Puszczy Bia-
łowieskiej. Ochrona Środowiska i Zasobów Naturalnych 40:
Malzahn E., Kwiatkowski W., Pierzgalski E. 2009. Przyroda nie-
ożywiona, w: Białowieski Park Narodowy. Poznać – Zrozu-
mieć – Zachować. (red. C. Okołów, M. Karaś, A. Bołbot) BPN,
Białowieża, 18–36. ISBN 9788387054687.
Matuszkiewicz J.M. 2007a. Zmiany w grądach, borach mieszanych
i łęgach jesionowo-olszowych puszczy Białowieskiej, w: Ge-
obotaniczne rozpoznanie tendencji rozwojowych zbiorowisk
leśnych w wybranych regionach Polski. (red. J.M. Matuszkie-
wicz). Instytut Geograi i Przestrzennego Zagospodarowania
PAN, Monograe 8, 197–289. ISBN 978-83-87954-78-0.
Matuszkiewicz J.M. 2007b. Ogólne kierunki zmian w zbiorowi-
skach leśnych polski, ich przyczyny oraz prognoza przyszłych
kierunków rozwojowych, w: Geobotaniczne rozpoznanie ten-
dencji rozwojowych zbiorowisk leśnych w wybranych regio-
nach Polski. (red. J.M. Matuszkiewicz). Instytut Geograi i
Przestrzennego Zagospodarowania PAN, Monograe 8: 670–
688. ISBN 978-83-87954-78-0.
Matwiejuk, A., Bohdan, A. 2011. New records of Plectocarpon
lichenum from Białowieża Forest (Poland). Herzogia 24: 381–
383. DOI 10.13158/heia.24.2.2011.381.
Matwiejuk A., Bohdan A. 2014. New sites of Bryoria capillaris
(Ach.) Brodo & D. Hawksw. (Ascolichenes, Parmeliaceae) in
the Polish part of the Białowieża Forest. Roczniki Akademii
Rolniczej w Poznaniu 390, Botanika Steciana 18(3): 181–185.
McKinney M.L., Lockwood J.L. 1999. Biotic homogenization: a
few winners replacing many losers in the next mass extinction.
Trends in Ecology and Evolution 14: 540–453.
Mikusińska A., Zawadzka B., Samojlik T., Jędrzejewska B., Mikus-
iński G. 2013. Quantifying landscape change during the last two
centuries in Białowieża Primeval Forest. Applied Vegetation
Science 16: 217–226. DOI 10.1111/j.1654-109X.2012.01220.x.
Naaf T., Kolk J. 2015. Colonization credit of post-agricultural forest
patches in NE Germany remains 130–230 years after reforesta-
tion. Biological Conservation 182: 155–163. DOI 10.1016/j.
Naaf T., Wulf M. 2010. Habitat specialists and generalists drive
homogenization and differentiation of temperate forest plant
communities at the regional scale. Biological Conservation
143: 848–855. DOI 10.1016/j.biocon.2009.12.027.
Nascimbene J., Thor G., Nimis P.L. 2013. Effects of forest man-
agement on epiphytic lichens in temperate deciduous forests
of Europe – A review. Forest Ecology and Management 298:
27–38. DOI 10.1016/j.foreco.2013.03.008.
Niemelä T. 2013. Polypores of the Białowieża Forest. Białowieski
Park Narodowy, Białowieża, 135 s. ISBN 978-83-87054-19-9.
Nespiak A. 1959. Studia nad udziałem grzybów kapeluszowych
w zespołach leśnych na terenie Białowieskiego Parku. Mono-
graphiae Bototanicae 8: 3–141.
Niklasson M., Zin E., Zielonka T., Feijen M., Korczyk A.F., Chur-
ski M., Samojlik T., Jędrzejewska B., Gutowski J.M., Brze-
ziecki B. 2010. 350-year tree-ring re record from Białowieża
Primeval Forest, Poland: implications for Central European
lowland re history. Journal of Ecology 98: 1319–1329. DOI
Okołów C. 2015. Materiały do oceny bioróżnorodności Puszczy Biało-
wieskiej. Nowe dla nauki gatunki organizmów opisane z Puszczy
Białowieskiej. Parki Narodowe i Rezerwaty Przyrody 34(3): 89–98.
Olden J.D., Poff N.L., Douglas M.R., Douglas M.E., Fausch K.D.
2004. Ecological and evolutionary consequences of biotic ho-
mogenization. Trends in Ecology and Evolution 19: 18–24.
Orłoś H. 1960. Badania nad funkcją ekologiczną grzybów z rodziny
Polyporaceae w różnych typach lasu Białowieskiego Parku Na-
rodowego. Prace Instytutu Badawczego Leśnictwa 193: 5–100.
Paluch R. 2009. Monitoring stanowisk granicznika płucnika (Lo-
baria pulmonaria L.) w lasach Nadleśnictwa Białowieża i za-
lecenia praktyczne dla jego ochrony. Leśne Prace Badawcze
321A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
Pawłowski J., Kubisz D., Mazur M. 2002. Coleoptera Chrząszcze,
w: Czerwona lista zwierząt ginących i zagrożonych w Polsce.
(red. Z. Głowaciński). Instytut Ochrony Przyrody Polskiej
Akademii Nauk, Kraków, 88–110. ISBN 83-901236-8-1.
Peterken G.F. 1974. A method for assessing woodland ora for
conservation using indicator species. Biological Conservation
6: 239–245. DOI 10.1016/0006-3207(74)90001-9.
Peterken G.F. 1996.Natural woodland. Ecology and conservation
in northern temperate regions. University Press, Cambridge –
New York – Melbourne, 540 s. ISBN 0521367929.
Pierzgalski E., Boczoń A., Tyszka J. 2002. Zmienność opadów i
położenia wód gruntowych w Białowieskim Parku Narodo-
wym. Kosmos 51(4): 415–425.
Pilát A. 1950. Contribution to the knowledge of the Hymenomy-
cetes of Białowieża virgin forest in Poland. Studia Botanica
Čechoslovaca 11: 145–173.
Podgórski T., Schmidt K., Kowalczyk R., Gulczyńska A. 2008.
Microhabitat selection by Eurasian lynx and its implications
for species conservation. Acta Theriologica 53: 97–110. DOI
Polski Komitet ds. UNESCO 2016a. http://www.unesco.pl/?id=290
Polski Komitet ds. UNESCO 2016b. http://www.unesco.pl/nauka/
Popławska. M. 2012. Porosty dębu szypułkowego Quercus robur
L. w Białowieskim Parku Narodowym. Praca magisterska wy-
konana w Zakładzie Botaniki Instytut Biologii Uniwersytetu w
Projekty ustaw 2006. Puszcza Białowieska. Dziedzictwo przyrod-
nicze i kulturowe. Projekty ustaw. https://www.mos.gov.pl/
Pucek Z. 2001a. Dryomys nitedula, w: Polska czerwona księga zwie-
rząt. Kręgowce. (red. Z. Głowaciński). Państwowe Wydawnic-
twa Rolnicze i Leśne, Warszawa, 77–79. ISBN 83-09-01735-9.
Pucek Z. 2001b. Sorex caecutiens, w: Polska czerwona księga zwie-
rząt. Kręgowce. (red. Z. Głowaciński). Państwowe Wydawnic-
twa Rolnicze i Leśne, Warszawa, 41–42. ISBN 83-09-01735-9.
Pucek Z., Jurczyszyn M. 2001. Glis glis, w: Polska czerwona księga
zwierząt. Kręgowce. (red. Z. Głowaciński). Państwowe Wydaw-
nictwa Rolnicze i Leśne, Warszawa, 79–81. ISBN 83-09-01735-9.
Pugacewicz E. 1997. Ptaki lęgowe Puszczy Białowieskiej. PTOP,
Białowieża. ISBN 83-903553-5-3.
Pykälä J. 2004. Effects of new forestry practices on rare epiphyt-
ic macrolichens. Conservation Biology 18: 831–838. DOI
Rachwald A., Ruczyński I. 2015. Common pipistrelle (Pipistrellus
pipistrellus Schreber, 1774) in the bat fauna of the Białowieża
Primeval Forest. Leśne Prace Badawcze 76(2): 180–183. DOI
Razowski J. (red.) 1990. Wykaz zwierząt Polski. T. 1. Wrocław
-Warszawa-Kraków, Zakład Narodowy im. Ossolińskich, Wyd.
PAN, 158 s.
Razowski J. (red.) 1991a. Wykaz zwierząt Polski. T. 2. Wrocław
-Warszawa-Kraków, Zakład Narodowy im. Ossolińskich, Wyd.
PAN, 342 s.
Razowski J. (red.) 1991b. Wykaz zwierząt Polski. T. 3. Kraków,
Krakowskie Wyd. Zool., 217 s.
Razowski J. (red.) 1997a. Wykaz zwierząt Polski. T. 4. Kraków, Wyd.
Inst. Syst. i Ewol. Zwierząt PAN, 303 s. ISBN 83-907187-0-7.
Razowski J. (red.) 1997b. Wykaz zwierząt Polski. Kraków, Wyd. Inst.
Syst. i Ewol. Zwierząt PAN, vol. 5, 260 s. ISBN 83-907187-1-5.
RDLP 2011. Program gospodarczo-ochronny Leśnego Kompleksu
Promocyjnego „Puszcza Białowieska” na lata 2012–2021. Re-
gionalna Dyrekcja Lasów Państwowych, Białystok.
Rose F. 1976. Lichenological indicators of age and environmental
continuity in woodlands, in: Lichenology: progress and prob-
lems (edit. D.H. Brown, D.L. Hawksworth, R. H. Bailey) Aca-
demic Press, London-New York, 279–307.
Rose F. 1992: Temperate forest management: its effects on bryo-
phyte and lichen oras and habitats, in: Bryophytes and lichens
in a changing environment. (edit. J.W. Bates, A. Farmer) Ox-
ford, Clarendon Press, 211–233. ISBN 0198542917.
Rose F., Coppins A.M. 2002. Site assessment of epiphytic habitats
using lichen indices, in: Monitoring with lichens – monitoring
lichens. (edit. P.L. Nimis, C. Scheidegger, P.A. Wolseley) Klu-
wer Academic Publisher, Dordrecht-Boston-London, 343–348.
Rowiński P. 2010. Puszcza Białowieska, w: Ostoje ptaków o zna-
czeniu międzynarodowym w Polsce. (red. T. Wilk, M. Jujka, J.
Krogulec, P. Chylarecki) OTOP, Marki. ISBN: 83-89830-02-7.
Rozporządzenie 2005. Rozporządzenie Wojewody Podlaskiego Nr
7/05 z dnia 25 lutego 2005r. w sprawie Obszaru Chronionego
Krajobrazu „Puszcza Białowieska”. Dz.U. Województwa Pod-
laskiego 54, poz. 720.
Rozporządzenie 2011. Rozporządzenie Ministra Środowiska z dnia
12 stycznia 2011 r. w sprawie obszarów specjalnej ochrony
ptaków. Dz.U. 25, poz. 133.
Rozporządzenie 2014a. Rozporządzenie Ministra Środowiska z
dnia 9 października 2014 r. w sprawie ochrony gatunkowej
grzybów. Dz.U. 2014, poz. 1408.
Rozporządzenie 2014b. Rozporządzenie Ministra Środowiska z dnia
7 listopada 2014 r. w sprawie ustanowienia planu ochrony dla
Białowieskiego Parku Narodowego. Dz.U. 2014, poz. 1735.
Ruczyński I., Bogdanowicz W. 2005. Roost cavity selec-
tion by Nyctalus noctula and N. leisleri (Vespertilioni-
dae, Chiroptera) in Białowieża Primeval Forest, eastern
Poland. Journal of Mammalogy 86: 921–930. DOI
Ruczyński I., Bogdanowicz W. 2008. Summer roost selection by
tree-dwelling bats Nyctalus noctula and N. leisleri: a mul-
ti-scale analysis. Journal of Mammalogy 89: 942–951. DOI
Ruczyński I., Ruczyńska I. 2000. Roosting sites of Leisler’s bat
Nyctalus leisleri in Białowieża Forest – preliminary results.
Myotis 37: 55–60.
Russo D., Cistrone L., Jones G., Mazzoleni S. 2004. Roost selec-
tion by barbastelle bats (Barbastella barbastellus, Chiroptera:
Vespertilionidae) in beech woodlands of central Italy: conse-
quences for conservation. Biological Conservation 117: 73–81.
Rydzak J. 1961. Tree lichens in the forest communities of the
Białowieża National Park. Annales Universitatis Mariae Curie
-Skłodowska, Sectio C 16(2): 17–47.
322 A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
Ryś A. 2007. Granicznik płucnik Lobaria pulmonaria i jego ochro-
na w Lasach Państwowych. Studia i Materiały Centrum Eduk-
acji Przyrodniczo-Leśnej 2/3(16): 288–302.
Samojlik T. 2010. Traditional utilisation of Białowieża Primeval
Forest (Poland) in the 15th–18th centuries. Landscape Archae-
ology and Ecology 8: 150–164.
Samojlik T., Fedotova A., Kuijper D.P. 2016. Transition from
traditional to modern forest management shaped the spatial
extent of cattle pasturing in Białowieża Primeval Forest in
the nineteenth and twentieth centuries. Ambio DOI 10.1007/
Samojlik T., Rotherham I., Jędrzejewska B. 2013. Quantifying
historic human impacts on forest environments: a case study
in Białowieża Forest, Poland. Environmental History 18(3):
576–602. DOI 10.1111/j.1654-109X.2012.01220.x.
Scheidegger C, Werth S. 2009. Conservation strategies for lichens:
insights from population biology. Fungal Biology Review
23(3): 55–66. DOI 10.1016/j.fbr.2009.10.003.
Schmidt K., Podgórski T., Kowalczyk R., Gulczyńska A. 2007.
O wymaganiach środowiskowych rysia eurazjatyckiego Lynx
lynx do bezpośredniego wykorzystania w aktywnej ochronie
gatunku w Polsce. Studia i Materiały Centrum Edukacji Przy-
rodniczo-Leśnej 9(2/3): 446–456.
Schmidt K., Jędrzejewski W., Theuerkauf J., Kowalczyk R.,
Okarma H., Jędrzejewska B. 2008. Reproductive behavior of
wild-living wolves Białowieża Primeval Forest (Poland). Jour-
nal of Ethology 26: 69–78.
SDF 2014. NATURA 2000 – STANDARDOWY FORMULARZ DA-
NYCH dla obszarów specjalnej ochrony (OSO), proponowanych
obszarów mających znaczenie dla Wspólnoty (pOZW), obszarów
mających znaczenie dla Wspólnoty (OZW) oraz specjalnych ob-
szarów ochrony (SOO). Puszcza Białowieska PLC200004. http://
Skirgiełło A. 1960. Wiosenne miseczniaki Białowieży. Mono-
graphiae Botanicae 10(2): 3–19.
Skirgiełło A. 1998. Macromycetes of oak-hornbeam forests in the
Białowieża National Park – monitoring studies. Acta Mycolo-
gica 33: 171–189.
Sokołowski A.W. 1995. Flora roślin naczyniowych Puszczy Bia-
łowieskiej, Białowieski Park Narodowy, Białowieża. ISBN
Sokołowski A.W. 2004. Lasy Puszczy Białowieskiej. Cen-
trum Informacyjne Lasów Państwowych, Warszawa. ISBN
Speight M.C.D. 1989. Saproxylic invertebrates and their conserva-
tion. Nature and Environment Ser. Strasbourg 42: 1–82.
Stachura K., Niedziałkowska M., Bartoń K. 2004. Różnorodność
ssaków leśnych, w: Eseje o ssakach Puszczy Białowieskiej.
(red. B. Jędrzejewska, J.M. Wójcik). Zakład Badania Ssaków
PAN, Białowieża, 13–24. ISBN 83-907521-2-3.
Szczepkowski A., Kujawa A., Karasiński D., Gierczyk B. 2008. Grzy-
by zgromadzone na XIV Wystawie Grzybów Puszczy Białowie-
skiej. Parki Narodowe i Rezerwaty Przyrody 27(4): 115–133.
Szczepkowski A., Kujawa A., Karasiński D., Gierczyk B. 2011.
XVII Wystawa Grzybów Puszczy Białowieskiej w Hajnówce.
Parki Narodowe i Rezerwaty Przyrody 30(3-4): 129–134.
Szujecki A. 2008. Puszcza Białowieska. Konikty wokół ochrony i
zarządzania. Centrum Informacyjne Lasów Państwowych.
Szujecki A. 2014. Sepedophilus wankowiczi (Pandellé, 1869) (Co-
leoptera: Staphylinidae) w Białowieskim Parku Narodowym.
Wiadomości Entomologiczne 33(2): 152.
Ściński M., Borowski Z. 2006. Home ranges, nest sites and po-
pulation dynamics of the forest dormouse Dryomys nitedula
(Pallas) in an oak-hornbeam forest: a live-trapping and radio
-tracking study. Polish Journal of Ecology 54: 391–396.
Ściński M., Borowski Z. 2008. Spatial organization of the fat dor-
mouse (Glis glis) in an oak-hornbeam forest during the mating
and post-mating season. Mammalian Biology 73: 119–127.
Thompson K., Bakker J.P., Bekker R. M. 1997. The soil seed bank of
North Western Europe: methodology, density and longevity. Cam-
bridge University Press, Cambridge, 276 s. ISBN 0 521 49519 9.
Tilman E., May R.M., Lehman C.L., Nowak M. 1994. Habitat de-
struction and the extinction debt. Nature 371: 65–66.
Tomiałojć L., Wesołowski T., Walankiewicz W. 1984. Breeding
bird community of a primeval temperate forest (Białowieża
National Park, Poland). Acta Ornithologica 20: 241–310.
Tomiałojć L., Wesołowski T. 2005. The avifauna of the Białowieża
Forest: a window into the past. British Birds 98: 174–193.
UNESCO 1984. Action plan for biosphere reserves, Nature and
Resources 20(4): 1–12.
van der Kooij J., Bangjord G., Obuch J., Carlsson B.-G., Hörnfeldt B.
2015. The range of the masked shrew (Sorex caecutiens Laxmann,
1788) extends to southern Scandinavia. Lutra 58(2): 119–126.
Vellend M., Verheyen K., Jacquemyn H., Kolb A., Van Calster H.,
Peterken G., Hermy M. 2006. Extinction debt of forest plants
persists for more than a century following habitat fragmenta-
tion. Ecology 87: 542–548. DOI 10.1890/05-1182.
Walankiewicz W. 2002. Nest predation as a limiting factor to the
breeding population size of the Collared Flycatcher Ficedula
albicollis in the Białowieża National Park (NE Poland). Acta
Ornithologica 37: 91–106. DOI 10.3161/068.037.0205.
Walankiewicz W. 2006. Czynniki ograniczające zagęszczenia lęgo-
we muchołówki białoszyjej Ficedula albicollis w pierwotnych
grądach Białowieskiego Parku Narodowego (z krytycznym
przeglądem wcześniejszych hipotez). Wydawnictwo Akademii
Podlaskiej. Rozprawa Naukowa nr 86.
Walankiewicz, W., Czeszczewik, D., Chylarecki, P. 2011. Dzięcioł
białogrzbiety Dendrocopos leucotos na obszarze Puszczy Bia-
łowieskiej w 2010 roku: rozmieszczenie, zmiany liczebności,
zagrożenia i perspektywy przetrwania populacji. Pracownia na
Rzecz Wszystkich Istot. Białystok-Siedlce-Warszawa.
Walankiewicz W., Czeszczewik D., Stański T., Sahel M., Ruczyński.
I. 2014. Tree cavity resources in spruce-pine managed and pro-
tected stands of the Białowieża Forest, Poland. Natural Areas
Journal 34: 423–428. DOI 10.3375/043.034.0404.
Wanat M. 1994. Ryjkowce (Coleoptera: Curculionoidea: Anthribi-
dae, Rhinomaceridae, Rhynchitidae, Attelabidae, Apionidae,
Curculionidae) Puszczy Białowieskiej. Polskie Pismo Ento-
mologiczne 63(1–2): 37–112.
Wanat M. 1999. Ryjkowce (Coleoptera: Curculionoidea bez Scoly-
tidae i Platypodidae) Puszczy Białowieskiej – charakterystyka
fauny. Parki Narodowe i Rezerwaty Przyrody 18(3): 25–47.
323A. Kujawa et al. / Leśne Prace Badawcze, 2016, Vol. 77 (4): 302–323
Wawrusiewicz A. 2011. Okres neolitu i wczesnej epoki brązu na
Podlasiu, stan i perspektywy badań, w: Na rubieży kultur.
Badania nad okresem neolitu i wczesną epoką brązu. (red. U.
Stankiewicz, A. Wawrusiewicz) Muzeum Podlaskie w Białym-
stoku, Białystok, 13–36. ISBN 978-83-87026-05-9.
Weiner J. 2016. Po co nam puszcze. Tygodnik Powszechny 27.06.-
Wesołowski T. 2005. Virtual conservation: How the European Union is
turning a blind eye to its vanishing primeval forests. Conservation
Biology 19: 1349–1358. DOI 10.1111/j.1523-1739.2005.00265.x.
Wesołowski T. 2007. Primeval conditions – What can we learn
from them? Ibis 149, Supplement 2: 64–77.
Wesołowski T., Czeszczewik D., Hebda G., Maziarz M., Mitrus
C., Rowiński P. 2015. 40 years of breeding bird community
dynamics in a primeval temperate forest (Białowieża National
Park, Poland). Acta Ornithologica 50: 95–120. DOI 10.3161/0
Wesołowski T., Fuller R.J. 2012. Spatial variation and tempo-
ral shifts in habitat use by birds at the European scale, in:
Birds and Habitat: relationships in Changing Landscapes
(edit. R.J. Fuller) Cambridge University Press, 63–92. ISBN
Wesołowski T., Kujawa A., Bobiec A., Bohdan A., Buchholz
L., Chylarecki P., Engel J., Falkowski M., Gutowski J.M.,
Jaroszewicz B., Nowak S., Orczewska A., Mysłajek R.W.,
Walankiewicz W. 2016. Spór o przyszłość Puszczy Biało-
wieskiej: mity i fakty. Głos w dyskusji. Chrońmy Przyrodę
Ojczystą 72(2): 83–99.
Wesołowski T., Rowiński P. 2006. Tree defoliation by winter moth
Operophtera brumata L. during an outbreak affected by struc-
ture of forest landscape. Forest Ecology and Management 221:
299–305. DOI 10.1016/j.foreco.2005.10.023.
Whigham D.F. 2004. Ecology of woodland herbs in temperate decidu-
ous forests. Annual Review of Ecology, Evolution, and Systematics
35: 583–621. DOI 10.1146/annurev.ecolsys.35.021103.105708.
Wolsan M., Okarma H. 2001. Lynx lynx, w: Polska czerwona
księga zwierząt. Kręgowce. (red. Z. Głowaciński) Państwo-
we Wydawnictwo Rolnicze i Leśne, Warszawa, 96–97. ISBN
Wołoszyn B.W. 2001. Nyctalus leisleri, w: Polska czerwona księga
zwierząt. Kręgowce. red. Z. Głowaciński) Państwowe Wydawnic-
two Rolnicze i Leśne, Warszawa: 58–59. ISBN 83-09-01735-9.
Zalewska A. 2012. Ecology of lichens of the Puszcza Borecka for-
est (NE Poland). W. Szafer Institute of Botany, Polish Acade-
my of Sciences, Kraków, 358 s. ISBN 978-83-62975-14-3.
Zalewska A., Bohdan A. 2012. New records of Lobaria amplissima
(Lobariaceae, Ascomycota) in Poland. Acta Mycologica 47(1):
97–108. DOI 10.5586/am.2012.012.
Zalewski A. 1997. Patterns of resting site use by pine marten Mar-
tes martes in Białowieża National Park (Poland). Acta Therio-
logica 42: 153–168.
Zarządzenie 1994. Zarządzenie nr 30 Dyrektora Generalnego
Lasów Państwowych z dnia 19 grudnia 1994 r. w sprawie Le-
śnych Kompleksów Promocyjnych (LKP). ZO-72-15/94
Zarządzenie 2015. Zarządzenie Regionalnego Dyrektora Ochrony
Środowiska w Białymstoku z dnia 6 listopada 2015 r. sprawie
ustanowienia planu zadań ochronnych dla obszaru Natura 2000
Puszcza Białowieska PLC200004. Dziennik Urzędowy Woje-
wództwa Podlaskiego, poz. 3600, Białystok.
Zbyryt A., Zawadzka D., Zawadzki G. 2014. Występowanie zająca
bielaka Lepus timidus w Polsce. Chrońmy Przyrodę Ojczystą
Zimny M. 2014. Późnoholoceńska historia roślinności Puszczy
Białowieskiej. Praca doktorska. Uniwersytet Gdański, Gdańsk.
All authors – the concept, assumptions, work system;
A.K. – coordination, writing, editing the whole manuscript;
A.K., A.O., M. B., M.F., J.M.G. – general chapters (Intro-
duction, Species Diversity in the Białowieża Forest – selec-
ted examples, Conclusions).
M.L. – history of the Białowieża Forest; M.F., A.O., A.K.
– nature conservation in the Białowieża Forest – current sta-
tus and threats; M.F. – evaluation of the conservation status
of Natura 2000 habitats in the Białowieża Forest and recom-
mendations on their protection; A.K. – macrofungi; A.Z.,
AB – lichens; A.O. – vascular ora; J.M.G., L.B. – insects;
P.Ch., W.W. – birds; S.N., R.W.M. – mammals.
Translated by: Bożena Kornatowska