The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park

Abstract

The results of palynological research presented here offer an insight into the longterm
dynamics of forest communities and the most important forest-forming species
in the area of the Białowieża National Park’s Strict Reserve, over the period of the last
2000 years. During this time, the Forest has developed under conditions in which climate
change occurred many times, and with levels of anthropogenic pressure also varying.
The most intensive human activity in the area of Strict Reserve took place in the Roman
Iron Age, as the Wielbark Culture tribes settled here, and then again in the 17th and
18th centuries, when a series of forest industries were permitted. Though material for
the study was mostly sampled at sites in close proximity to many archaeological traces
of past human activity, the pollen diagrams indicate only a limited infl uence of that
activity on the natural environment. The regeneration of forest communities took place
here with no more major involvement of pioneer species, and thus led to a return to
the situation prior to disturbances, with this attesting to maintained continuity of forest
habitats. What is noteworthy are the relatively small proportions (if we consider
the period) of indicators of cultivation. This is again suggesting local farming on just
a very limited scale what is in accordance with historical documents reporting on about
400-years restrictions imposed on settlement development in Białowieża Forest. This was presumably the main factor allowing ecosystems in the Forest to remain in a condition
allowing them to be identifi ed as “natural” in line with the classifi cation of the international
group of experts dealing with forest terminology (Buchwald 2005).

Full text

The Forests of the Strict Reserve of Białowieża National Park
The Forests
The Forests
of the Strict
of the Strict
Reserve
Reserve
ofBiałowieża
ofBiałowieża
National Park
National Park
It has been areal pleasure tobecome acquainted with this new scientifi c study concerning Białowieża Forest. Alarge
group ofscientists representing the Faculty ofForestry ofWarsaw University ofLife Sciences (SGGW), the University
ofGdańsk and Białowieża National Park here bring toaclose agreat many years ofwork devoted tothe forest inBNP’s
Strict Reserve. Specifi cally, this publication sets out topresent the story behind the creation ofone ofPoland’s oldest
protected areas, while also offering analysis ofthe dynamics that have characterised the development ofthevarious
tree species present here, down the ages. Aninteresting methodology founded upon anetworkofcircular permanent
plots has sustained adetailed characterisation ofthe Reserve’s stands from the point ofview ofdead trees, while
the information gathered onthe permanent plots has also allowed for aforecast ofstand development over the
next 20 years. Thus, inwhat ismulti-stranded, fascinating work, welearn ofthe developmental phases stands pass
through and the processes ongoing inthem, while the results ofhistorical and recent inventories oftrees achieving
monumental dimensions are also presented. Studies and analyses carried out support assessments ofthecondition
forest communities embraced bythe
Natura 2000
programme are tobefound in.
The Forests ofthe Strict Reserve ofBiałowieża National Park
isavery valuable new scientifi c work that does much
toincrease our knowledge ofthe functioning ofwhat are forest communities entirely unique anywhere inEurope.
Prof. Tomasz Borecki
Białowieża Forest is a “one-off”, whose exceptional nature is most fully encapsulated in its “Strict Reserve”. This
book characterises that area’s tree stands indetail, with information also offered on their history – all the way back
topalaeobotanical reconstructions ofchanges inplant cover taking place during the Holocene. Inthe 1930s, Prof.Tadeusz
Włoczewski ofwhat istoday Warsaw University ofLife Sciences (SGGW) wisely founded permanent plots bywhich stand
composition and structure could bestudied along agradient ofsite-types from alder swamp forest through todry mixed/
coniferous forest. Thankfully, the measurement data for the Strict Reserve resembled the Reserve itself insurviving the
turbulent times ofWorld War II;and from the 1950s on,asuccessor generation toProf. Włoczewski pressed onwith and
augmented his work. The data series become more precious with each passing decade, leaving this book inaposition
toset current research results against 80-year measurement and observation series. This ensures that wemay now truly
look atBiałowieża Forest interms ofboth the processes that formed it,and those that goonshaping the composition
and structure ofunmanaged stands largely unaffected byhuman activity.
Prof. Jerzy Szwagrzyk
This publication was prepared within the frames
oftheproject
Fostering management of the Białowieża
National Park through harmonization of environmental data
co-fi nanced by Iceland, Liechtenstein and Norway
under the European Economic Area Financial Mechanism
ISBN 978-83-64513-15-2

The Forests
oftheStrict Reserve
ofBiałowieża
NationalPark
Edited by
Andrzej Keczyński


Authors:
K B, L B, B B, W B, S D,
L G, A G, A K, M L, A
P, A S, H S, J Z, MZ
Reviewers:
T B, Professor, Department of Forest Management Planning and Forest Economics,
Warsaw University of Life Sciences
J S, Professor, Department of Forest Biodiversity, University of Agriculture inKraków
Translator:
J R. A. R, PhD
Scientifi c editor:
A K, MSc, Eng.
Proofreading:
L H
Cover and typographic layout design:
W K
Typesetting and text make-up:
W K
Cover photography:
A K
Front: oaks in the sedge-dominated form of oak-lime-hornbeam forest (
Tilio-Carpinetum
caricetosum remotae
).
Back (top, from left to right): infl orescence of wych elm
Ulmus glabra
; multiple fruits of common
aspen
Populus tremula
; new generation of spruce living on dead spruces; Norway maple
Acer
platanoides
; (bottom, from left to right): butts of spruce and oak; bark of black alder
Alnus
glutinosa
; trunk and tree crown of oak
Quercus robur
; lying after hurricane dead trees in the
BNP Strict Reserve.
Photography edition:
M B
Illustrations of trees:
J I
Print preparation:
P K, PRE-TEKST, www.pre-tekst.com
Publisher:
Białowieski Park Narodowy
Park Pałacowy 11
17-230 Białowieża
www.bpn.com.pl
Contractor:
Wydawnictwo DRAGON Sp. zo.o.
© Copyright by Białowieski Park Narodowy 2017
All rights reserved
ISBN 978-83-64513-15-2
The project
Fostering management of the Białowieża National Park through harmonization
of environmental data
is partially funded by the EEA grants from Iceland, Liechtenstein
andNorway totalling 2 565 781 zł


Andrzej Keczyński
Introduction 5
Andrzej Keczyński
General description of the Strict Reserve of Białowieża National Park 9
Marcelina Zimny, Małgorzata Latałowa, Anna Pędziszewska
The Late-Holocene history of forests in the Strict Reserve
of Białowieża National Park 29
Jacek Zajączkowski, Bogdan Brzeziecki, AndrzejKeczyński
Living and dead (standing and lying) trees as major components of stands 61
Bogdan Brzeziecki
Long-term tree andstand dynamics 93
Kamil Bielak, Bogdan Brzeziecki
Current structural differentiation and forecast ofdevelopment
of the main woodland types 127
Stanisław Drozdowski, Bogdan Brzeziecki, Andrzej Keczyński
The dynamics to the different developmental phases 159
Stanisław Drozdowski, Włodzimierz Buraczyk, Bogdan Brzeziecki
The process of thenatural regeneration of forest 183
Andrzej Grzywacz, Andrzej Keczyński, Andrzej Szczepkowski, Kamil Bielak,
Stanisław Drozdowski, Leszek Bolibok, Bogdan Brzeziecki
Trees of monumental sizes 213
Bogdan Brzeziecki, Stanisław Drozdowski, Kamil Bielak,
Włodzimierz Buraczyk, Henryk Szeligowski, Leszek Gawron
Allometry ofthelargest trees 247
Andrzej Keczyński
Monitoring
Natura2000
forest habitat types 265
Bogdan Brzeziecki, Andrzej Keczyński, Jacek Zajączkowski
The permanent study plots 277
List of tables 294
List of fi gures 296
List of photographs 300
Authors 304
Contents

 A. Keczyński

General description ofthe Strict Reserve of Białowieża NationalPark
Marcelina Zimny, Małgorzata Latałowa,
Anna Pędziszewska
The Late-Holocene
history of forests
inthe Strict Reserve
of Białowieża
NationalPark
Illustration by J.Iwacik

 M. Zimny, M. Latałowa, A. Pędziszewska
The Late-Holocene
history of forests
inthe Strict Reserve
of Białowieża
NationalPark
Introduction
Trees are longlived organisms forming ecological systems that can often stay in dynamic
equilibrium for hundreds of years. However, over millennia, theconditions under which
thespecies building a forest community compete are infl uenced by different factors
like changes of climate, as well as such shorter-term natural events as fl oods, gales and
fi res, and mass outbreaks of insect pests and pathogens. Since themid-Holocene at
least, large areas of Europe have also been much infl uenced by human activity.
Depending on their intensity, frequency of occurrence and spatial scale of impact,
these factors have led over time to reversible processes of destruction and regen-
eration, in thecourse of which dominant species have changed places periodically
(F 1988) or to permanent remodelling of forest ecosystems (cf.B and
S 2014). However, given the length of time they take, these processes evade
direct scientifi c observation. Inthe relatively few cases in which research on perma-
nent plots has extended into periods measured in decades this is till only long enough
to supply data regarding certain short stages in thedevelopment of forests. Contem-
porary monitoring of forest habitats, while informative, has even less to offer when
change measured in centuries is involved. Thus, for aninsight into long-term changes
in thedynamics of forest communities, natural archives such as biogenic sediments in
lakes and peatlands should be analysed.
Regarded as “the best-preserved broadleaved and mixed/broadleaved forest complex
anywhere in theEuropean Lowlands” (F 1986; A 2014), theBiałowieża For-
est represents aunique environment enabling to study natural processes ongoing in
forest ecosystems (W
etal.
2016). Particularly priceless in this wider area is

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
thepart falling within today’s Strict Reserve, which has enjoyed protection for almost
100years now. This area has perfect documentation as regards thepresent-day fl ora
and forest communities (
i.a.
F 1986; S 1995, 2004; C 2008;
B
et al.
2011; B 2012) as well as acertain amount of archaeological and
historical data indicating local human activity (
i.a.
S
etal.
2013a, b). However,
unless thepalaeoecological record of thehistory of vegetation is concerned, no rel-
evant assessment can be made of either theecological impacts of former human
activity, or thenatural long-term dynamics of theforest-forming species; this kind
of knowledge offers for both tracing what has happened in thepast and considering
potential future changes (cf.B and S 2014).
The aim of this study has thus been to present key data on thelast 2000years of
history of forest communities in thearea of theStrict Reserve of BNP, by reference to
theresults of pollen analysis of sediments collected at six sites within theReserve,
plus two more located in its immediate vicinity.
Coniferous bog
forest
Vaccinio
uliginosi-Pinetum
in theBNP Strict
Reserve
Photograph by
A.Keczyński

 M. Zimny, M. Latałowa, A. Pędziszewska
Material and methods used for thepalaeoecological
reconstruction
The sites sampled for palaeoecological material are small, shallow peatbogs within
theforest, which differ in size, thecontemporary vegetation cover and thehabitats
present in thesurrounding areas (Fig.2). Sediment profi les collected from these bogs
(see photograph p. 34) were between 50and 110 cm in depth. The sediment cores
were transferred to the palynological unit at the Laboratory of Palaeoecology and
Archaeobotany (Department of Plant Ecology) of theUniversity of Gdańsk, where –
after detailed description – they were divided up into slices 0.5cm thick, before sup-
plying 0.5cm3samples for pollen analysis. Thesamples went through amulti-stage
laboratory procedure (F and I 1989), with theaim being to obtain material
for analysis that is as pure as possible.
Fig. 2. Locations
ofpalaeoecological
sites against
the background
ofamap showing
the distribution
ofplant communities
drawn upby
KWIATKOWSKI and
GAJKA (2009);
site symbols
refl ect the forest
compartment they
were includedin;
the MC1/256 site
was worked onby
MITCHELL and COLE
(1998)
Illustration by
M.Zimny palaeoecological sites

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
Palaeoecological
sites
Photograph by
M.Zimny
BIA/314D BIA/340G
BIA/369F BIA/317C
BIA/318C BIA/287D
BIA/131C BIA/161A
Microscopic analysis in thedifferent profi les was carried out to astratigraphic resolu-
tion of 0.5or 1cm, with identifi cations and counts being made of about 1000grains
of arboreal pollen (AP) plus thepollen of herbs – non-arboreal pollen (NAP) and other
microfossils comprising remains of organisms of different groups preserved in thepal-
ynological preparations, with these including algae, Cyanobacteria, fungi and rhizo-
pods (i.e. non-pollen palynomorphs or NPPs;  G 2001). Charcol microparticles

 M. Zimny, M. Latałowa, A. Pędziszewska
were counted as well (Tinner and Hu2003). Full data on this are included in separate
works (Z 2014; L
etal.
,
in preparation
; Z
etal.
,
in preparation
).
To determine theages of successive stages in thehistory of theforest, and assess
their durations, plant macrofossils were selected from different horizons in thepro-
fi les, these then serving as material for radiocarbon dating by Accelerator Mass Spec-
trometry
(AMS) at the Poznań Radiocarbon Laboratory. Obtained in this way were
39radiocarbon dates which, after appropriate calibration (using theOxCal program,
version 4.2: B R and L 2013; calibration curve from R
etal.
2013),
provided abasis for thechronological determinations. Anattempt was also made to
determine theages of the upper parts of theprofi les using the(210Pb) lead method
(O 1986; A 2001). Thedating was done at theDepartment of Quaternary
Fieldwork–
collecting peat
profi les for
palaeoecological
research
Photograph by
M.Zimny

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
Geology and Geomorphology of theUniversity of Gdańsk. Thefull basis for thechro-
nology established for palaeoecological events in thehistory of the BNP forests is
discussed in aseparate paper (L
etal.
,
in preparation
).
Here we present selected data that have been depicted in theform of three simpli-
fi ed pollen diagrams, i.e. two for sites BIA/314D (Fig.3) and BIA/369F (Fig.4) situ-
ated within theBNP Strict Reserve, and one for site BIA/131C (Fig.5), which is just
within thepresent (northern) boundary of BNP. Maps are also included (inFigs. 6–12)
to illustrate theproportions of thepollen of certain arboreal and non-arboreal taxa
in successive periods. Inthis case we used thedata of all of thesites studied by us
within theboundaries of today’s BNP, as well as one of thesites analysed by M-
 and C (1998) (source:
European Pollen Database:
www.cricyt.edu.ar/paleo/epd/
epd_main.html) whose chronology was corrected on thebasis of our data (L
etal.
,
in preparation
). Thepercentages accounted for thepollen of different taxa –
as presented on both diagrams and maps – refer to apollen sum of all arboreal and
non-arboreal taxa (AP + NAP = 100%), albeit with local taxa representative of wet-
lands excluded.
This study makes use of theterm “anthropogenic pollen (orpalynological) indicators”.
Inline with thecriteria proposed by B (1981), these include thepollen of cultivated
plants, as well as those spreading in thedifferent habitats shaped by human activity
(cf.L 2003, 2007). Atotal for pollen in this category can be treated as apaly-
nological indicator of anthropopressure.
The main stages of theHolocene history
ofvegetationin theBiałowieża Forest
Just as the interpretation of contemporary processes ongoing in plant cover is not
proper without alook back at long-term dynamics decades or even centuries into
thepast (B and S 2014), so also aproper appreciation of processes taking
place over thelast 2000years in thearea of what is today theStrict Reserve of BNP
requires alook even further back into thepast. Thelast 2000years represent just
themost recent stage to thehistory of vegetation in theHolocene, while theStrict
Reserve is just asmall part of aforest complex with avery specifi c history. Inthis
regard, it should be recalled that there is aseveral-thousand-year history to all of
thepresent-day forest communities in thearea under study, with thesingle exeption
of those whose ecology and physiognomy is shaped by amarked presence of Norway
spruce (L
etal.
2016).

 M. Zimny, M. Latałowa, A. Pędziszewska
Fig. 3. Simplifi ed
pollen diagram for
the site BIA/314D
profi le
Illustration by
M.Zimny
Lithology:
1–cotton-grass peat;
2–cotton-grass and bog-
moss peat; 3–bog-moss
peat
chronology
coprophilous fungi
Neurospora
Gelasinospora
Melampyrum
Pteridium aquilinum
Calluna vulgaris
ruderal weeds
Artemisia
segetal weeds
Centaurea cyanus
Rumex acetosella
-t.
cultivated plants
Fagopyrum
Secale cereale
Filipendula
Poaceae
Tilia cordata
Quercus
Acer
Fraxinus excelsior
Alnus
Populus
lithology
Pinus sylvestris
Triticum-
t.
meadow plants
Plantago lanceolata
anthropogenic indicators
Carpinus betulus
Corylus avellana
Ulmus
Picea abies
Salix
depth [cm]
Betula alba-
t.
charcoal > 40 m
20th
century
3
0
2
4
6
8
10
12
14
16
18
20
22
24
26
28
30
32
34
36
38
40
42
44
46
48
50
52
54
56
58
60 50 30 5 5 20 20 5 5 5 10 20 20 10 10 10 5 5 5 5 5 5 5 5 5 5 5 10 5 5 5 10 5 5 10 [%]
%%%%
‰‰‰‰
2
1
19th
century
17th–18th
century
9th–13th
century
6th–8th
century
1st–5th
century
HIATUS
HIATUS
BIA/314D

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
Fig. 4. Simplifi ed
pollen diagram for
the site BIA/369F
profi le
Illustration by
M.Zimny
Lithology:
1–sand; 2–herbaceous
peat with an admixture of
detritus gyttja; 3–sedge
and bog-moss peat
19th
century
3
6
8
10
12
14
16
18
20
22
24
26
28
30
32
34
36
38
40
42
45 30 20 10 10 20 20 5 5 5 10 30 30 10 10 10 5 10 10 5 5 5 10 5 5 5 5 10 5 5 5 10 5 5 10 [%]
%%%% ‰‰‰‰
2
1
18th
century
17th
century
BIA/369F
chronology
coprophilous fungi
charcoal > 40 m
Pteridium aquilinum
ruderal weeds
segetal weeds
Rumex acetosella
-t.
Fagopyrum
Secale cereale
Filipendula
Poaceae
Tilia cordata
Quercus
Acer
Fraxinus excelsior
Alnus
Populus
Pinus sylvestris
lithology
Anthoceros punctatus
Neurospora
Melampyrum
Calluna vulgaris
Artemisia
Centaurea cyanus
cultivated plants
Triticum-
t.
meadow plants
Plantago lanceolata
anthropogenic indicators
Carpinus betulus
Corylus avellana
Ulmus
Picea abies
Salix
Betula alba-
t.
depth [cm]

 M. Zimny, M. Latałowa, A. Pędziszewska
Fig. 5. Simplifi ed
pollen diagram for
the site BIA/131C
profi le
Illustration by
M.Zimny
Lithology:
1–cotton-grass and bog-
moss peat; 2–bog-moss
peat
BIA/131C
chronology
Neurospora
charcoal > 40 m
Pteridium aquilinum
ruderal weeds
segetal weeds
Rumex acetosella
-t.
Fagopyrum
Secale cereale
Filipendula
Poaceae
Tilia cordata
Quercus
Acer
Fraxinus excelsior
Alnus
Populus
lithology
Pinus sylvestris
coprophilous fungi
Gelasinospora
Melampyrum
Calluna vulgaris
Artemisia
Centaurea cyanus
Triticum-
t.
meadow plants
Plantago lanceolata
anthropogenic indicators
Carpinus betulus
Corylus avellana
Ulmus
Picea abies
Salix
depth [cm]
Betula alba-
t.
cultivated plants
0
2
4
6
8
10
12
14
16
18
20
22
24
26
28
30
32
34
36
38
40
42
44
46
48
50
52
54
56
58
60
62
64
%%% %
‰‰‰ ‰
2
1
50 30 5 5 30 20 5 5 5 10 10 10 5 10 10 5 5 5 5 5 5 10 5 5 5 5 5 5 5 5 20 5 5 5 [%]
20th
century
19th
century
18th
century
17th
century
HIATUS
5th–13th
century
1st–4th
century

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
The results of pollen analyses of thepeatbogs at Dziedzinka and Kletno (inthe south-
ern part of theStrict Reserve), done by M
etal.
(2009), as well as theCzerlon site
(L
etal.
2016) located just 6km to thesouth-west of theReserve boundary,
show that at thebeginning of theHolocene (i.e. some 9750–8800years BC) birch
forest with anadmixture of Scots pine and aspen dominated in thelandscape. From
about 9300BC elm was also present. Animportant element of theplant cover was
ascrub in which willows and shrub and dwarf-shrub birches were present.
It was around 8800years BC that pine largely displaced birch in thearea, becoming
themain component of theforest communities and hazel also appeared as amajor
species. Inthe period around 7300–7100BC amulti-species broadleaved forest was
already fully-developed. Its main components were oak, hazel, small-leaved lime, elm
and ash, with admixture of Norway maple. Spread across wet habitats were commu-
nities in which black alder prevailed.
A further radical restructuring of forest cover began about 1800 years BC, when
thehazel that had been represented so abundantly previously gave way to hornbeam.
Theforest that developed was thus of theoak-lime-hornbeam type, and did indeed
include large shares of oak and hornbeam, albeit amore limited representation of
lime than previously. Thelatter species resembled elm and ash in achieving optimal
status during theMiddle Holocene. Astill-important spatial role was played by swamp
and riverine communities with alder, even if that role was coming to be markedly
limited by around 2000years ago. Furthermore, through theentire Holocene, patches
of poorer soils continued to support forests in which Scots pine played amajor role.
At thesites, thesections of profi les dated to theMiddle Holocene are characterised
by thepresence of limited amounts of spruce pollen. However, this is not enough of
apremise to state unambiguously that this conifer was present locally in theperiod.
Palynological data from theCzerlon site suggest theoccurrence of alimited admixture
of this species in theforests about 4000BC, as well as afi rst more marked increase in
population density as recently as in themiddle of thefi rst millennium AD.
In line with palynological criteria, thefi rst traces of human activity in theCzerlon profi le
are to be noted in samples dating from between 2500and 1800years BC. Single pollen
grains of ribwort plantain (
P. lanceolata
), which is regarded as anindicator of grazing and
primitive agriculture (B 1981), presumably indicate local presence of aLate Neolithic
tribe, most likely representing theCorded Ware Culture which left themost numerous
archaeological traces in thePodlasie region in theperiod under concern (W
2011). Further, weak episodes of settlement are refl ected in asection of thediagram
dated to around 1200–650years BC (i.e. theLate Bronze Age and Early Iron Age).

 M. Zimny, M. Latałowa, A. Pędziszewska
Fig. 6. Mean
percentage
proportions
of
Carpinus
and
Quercus
pollen
atsites inBNP,
inconsecutive
periods; blue dots
denote alack ofdata
for the given period
Illustration by
M.Zimny
Carpinus
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
Quercus
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
<2% 2–5% 5.1–10% 10.1–15% >15%

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
Fig. 7. Mean
percentage
proportions of
Tilia
and
Corylus
pollen
atsites inBNP,
inconsecutive
periods; blue dots
denote alack ofdata
for the given period
Illustration by
M.Zimny
Tilia
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
Corylus
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
<0.1% 0.1–0.5% 0.51–1% 1.1–2% >2%

 M. Zimny, M. Latałowa, A. Pędziszewska
Fig. 8. Mean
percentage
proportions of
Ulmus
and
Fraxinus
pollen
atsites inBNP,
inconsecutive
periods; blue dots
denote alack ofdata
for the given period
Illustration by
M.Zimny
Ulmus
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
Fraxinus
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
<0.1% 0.1–0.5% 0.51–1% 1.1–2% >2%

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
Fig. 9. Mean
percentage
proportions of
Picea
and
Alnus
pollen
atsites inBNP
inconsecutive
periods; blue dots
denote alack ofdata
for the given period
Illustration by
M.Zimny
Picea
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
Alnus
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
<2% 2–5% 5.1–10% 10.1–15% >15%

 M. Zimny, M. Latałowa, A. Pędziszewska
Fig. 10. Mean
percentage
proportions of
Pinus
pollen aswell as
theratio ofarboreal
tonon-arboreal
pollen (AP/NAP)
atsites inBNP,
inconsecutive
periods; blue dots
denote alack ofdata
for the given period
Illustration by
M.Zimny
Pinus
1st–5th century 6th–8th century 9th–13th century
AP/NAP
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
AP NAP
<20% 20–30% 31–40% 41–50% >50%
17th–18th century 19th century 20th century

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
Fig. 11. Mean
percentage
proportions
of thepollen
ofanthropogenic
indicator species
and cereals
atsites inBNP,
inconsecutive
periods; blue dots
denote alack ofdata
for the given period
Illustration by
M.Zimny
Anthropogenic indicators
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
Cereals
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
<0.5%0% 0.5–1% 1.1–2% 2.1–5% >5%

 M. Zimny, M. Latałowa, A. Pędziszewska
Fig. 12. Mean
percentage
proportions
of
Artemisia
and
Calluna
pollen
atsites inBNP,
inconsecutive
periods; blue dots
denote alack ofdata
for the given period
Illustration by
M.Zimny
Artemisia
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
Calluna
1st–5th century 6th–8th century 9th–13th century
17th–18th century 19th century 20th century
<0.5%0% 0.5–1% 1.1–2% 2.1–5% >5%

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
The last two thousand years in thehistory
oftheforests in and around theStrict Reserve
The 1st to 5th centuries
Around 2000years ago, thekey components of broadleaved forest occuring in thearea
included pedunculate oak (
Quercus
robur
) and hornbeam (
Carpinus betulus
). Present as
major admixture species were (above all) small-leaved lime (
Tilia cordata
) and hazel
(
Corylus avellana
). Asthey do today, these species formed anoak-lime-hornbeam forest
community. More widespread than today were patches of alder carr, as well as riparian
forests dominated by alder (
Alnus glutinosa
), with anadmixture of ash (
Fraxinus excel-
sior
) and elm (
Ulmus
sp.). Onlighter soils therole of Scots pine (
Pinus sylvestris
) was
considerable. Inthe local stands presence of spruce (
Picea abies
) was still rather limited.
From thepalynological point of view, thesections of theprofi les dated to this period
are characterised by wide-amplitude fl uctuations of thepollen curves for broadleaved
trees, and high proportions of non-arboreal pollen in the4.5–15.7% range. Thelat-
ter include crop plants and species typical for anthropogenic habitats (Figs. 3and 5),
thereby attesting to thepresence of patches of non-forest communities, whose spread
should be linked with anearby settlement. Local development of settlements is in fact
confi rmed archaeologically as relating fi rst to theStroked Ware Culture, and then to
theWielbark Culture (K
etal.
2008; S
etal.
2013b). Three sites of
theformer have been discovered within between 2and 6km of theStrict Reserve’s
boundaries, while two of thefour known sites for theWielbark Culture are situated
directly within thearea of theReserve (L
etal.
2015).
At the time in question, the population dwelling here made use of burning tech-
niques. Palynological samples reveal many charcoal microparticles, as well as spores
of
Gelasinospora
fungi that habitually develop on burnt organic matter ( G and
A 2006), and pollen and spores from light-demanding plants frequent in fi re-af-
fected habitats –heather (
Calluna vulgaris
), cow-wheat (
Melampyrum
sp.) and bracken
(
Pteridium aquilinum
). Destruction in multispecies broadleaved forest habitats also
allowed for ashort-term expansion by light-demanding tree species as shown by
higher proportions of birch and pine pollen.
Land cleared from forest was made use of in farming. Samples include thepollen of
crop plants, as well as of plants typical for grazed areas. However, its proportions are
relatively low. Aparticularly good refl ection of thesettlement phase of theWielbark
Culture was obtained at sites BIA/314D (Fig.3), BIA/340G, and especially site BIA/318C

 M. Zimny, M. Latałowa, A. Pędziszewska
located close to remnants of aburial ground (Wielka Kletna, compartment 345A) – which
was probably in use between the2nd and 5th centuries AD (K
etal.
2008).
Barley and wheat (
Hordeum
-t.,
Triticum
-t.) were under cultivation, as well as rye (
Secale
cereale
) and hemp (
Cannabis sativa
), whose local use gained further confi rmation when
seeds of thespecies were found among macroscopic remains in theprofi le at BIA/318C.
A key further element to theland use lay in theraising of livestock, which grazed and
browsed in thehabitats created once patches of oak-lime-hornbeam forest had been
removed (with this being attested to by thepresence of pollen of ribwort plantain), as
well as in river valleys where theremoval of riparian forests allowed for thespread
of wet meadows featuring meadowsweet (
Filipendula ulmaria
). Ahypothesis regard-
ing opportunity for forest grazing to take place in forests thinning with theuse of
fi re is attested to by theregular presence of thepollen of heather and sheep’s-bit
scabious (
Jasione montana
). High level of disturbance in forest habitats (asattested
to by thepollen diagrams), in thecontext of arelatively limited proportions of crop
plants pollen, might be explained at least in part by metallurgical activity, of which
there are indeed archaeological traces in theform of theremains bloomer furnaces
and slag heaps known from theBiałowieża Forest (J 1939; S 2007).
Theextraction of bog iron no doubt led to thedestruction of riparian habitats, while
charcoal-burning obviously required that large amounts of wood be procured.
The 6th to 13th centuries
The end of the5th century brought abreakdown of Wielbark Culture settlements along
theprocesses of theGerman tribes migratory movements (M 2013). Asettle-
ment crisis also took place in thearea under study, with theeffect being aregeneration
of forest on land that had earlier been in use – as is seen in anincrease in theshare of
tree pollen (Fig.10), especially that of hornbeam (Fig.6), as well as aminimal propor-
tions of pollen indicative of anthropogenic activity (Fig.11).
First and foremost it was populations of hornbeam and oak that rebuilt, with other
components of broadleaved forest only doing so to amore limited extent. Therole
played by spruce also increased at this stage, as favoured by theearlier disturbances
of forest habitat, as well as conditions of acool and wet climate favourable to this
species (B
etal.
2011). Theissue under discussion remains that of thepos-
sible persistence of small settlements in thearea of the Białowieża Forest during
themigration period. This idea is suggested, not only by theconstant presence of
charcoal microparticles and high frequencies of heather pollen (Fig.12), but also by

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
thepresence of small amounts of pollen from fi rst-order indicators of anthropogenic
activity, especially cereals and ribwort plantain (Figs. 3, 5and 11) (L
etal.
2015,
2016; Z
etal.
,
in preparation
).
Major changes in forest cover began between the9th and 10th centuries, as arefl ection of
aregression of alder (Fig.9), as well as – to alesser extent – ash, oak and hornbeam. This
was also thetime at which spruce started to expand. Themain impulse behind these pro-
cesses would seem to have been thechange of climate described as “Mediaeval Climate
Anomaly” (
i.a.
B
etal.
2011), which would generate afall in water table in swampy
areas of theBiałowieża Forest (L
etal.
,
in preparation
), and then theencroach-
ment of spruce into habitats previously occupied by alder carr. This may be presumed to
be theperiod in which thespruce swamp forest community
Sphagno girgensohnii-Picee-
tum
fi rst began to develop within theBiałowieża Forest (L
etal.
2016).
Thus far, the period of theEarly Middle Ages is represented by only limited archaeo-
logical traces of settlement in theBiałowieża Forest, with acomplete lack of artefacts
in thearea of theStrict Reserve (K
etal.
2005, 2011; S
etal.
2013b;
L
etal.
2015). Palynological data only point to aminor increase in thepresence
of pollen indicating human activity for this period (Fig.11). This suggests that anthropo-
genic factors were only of secondary importance at this stage of thehistory of theforest
in theStrict Reserve, including also where theexpansion of spruce was concerned.
The results of pollen analysis indicate that economic activity in this area was also limited
in thelater phase of themedieval period. This is above all true of farming, with palyno-
logical samples only sporadically revealing thepresence of pollen from cereals. Pollen
of plants indicative of pastures are also infrequent. Incontrast, theconstant presence
of charcoal microparticles, as well as heather pollen and bracken spores, all attest to
disturbances in forest habitats, presumably refl ecting low-scale use of forest for animal
breeding, as well as to meet other local needs involving charcoal and potash-burning,
as well as thesmelting of bog iron. Inthis case also, palynological data are in line with
earlier archaeological and historical fi ndings, which point to settlement on only alimited
scale in theBiałowieża Forest up to the14th century, at least (S
etal.
2013a).
The 14th to 16th centuries
Due to disturbances of the sediments, none of the sites has retained more than
afragmentary depiction of theenvironmental history in theLate Middle Ages. Pol-
len diagrams do not therefore offer abasis for any more exhaustive characterisation

 M. Zimny, M. Latałowa, A. Pędziszewska
of changes taking place in thevegetation, most especially in the14th–15th centuries,
and partially also the16th century (L
etal.
,
in preparation
). Notwithstanding
thelimits of thepalaeoecological data, certain conclusions about potential changes
in forest cover may be drawn from historical sources describing settlement and uti-
lisation of thearea of theForest in this period. Upto the15th century, and hence to
thetime when regulations limiting theuse of theForest imposed by theRoyal admin-
istration came into force (S
etal.
2013b), thescale of local settlement was pre-
sumably limited, with earlier forms of utilisation of forest resources being maintained.
From the15th century onwards, theBiałowieża Forest came under protection, so that
hunting land might be available for theGrand Dukes of Lithuania and Kings of Poland.
Hunting manors and hunting enclosures – of which one (ofWielka Kletna) was located
in anarea that today forms part of theReserve – operated for this purpose (S
etal.
2013b). Inthe 15th–17th centuries, management of theForest revolved around
Royal hunts, though there were afew areas of meadow on which hay was collected,
areas with beehives and various removals of raw materials from theforest to meet
theneeds of those in theKing’s service. Dating to the16th century is small-scale pot-
ash-burning and tar-making (W 1984; S
etal.
2013a). However, thevery
limited scale of settlement (S
etal.
2013b), and severe restrictions on access to
ensure protection of theresources of theForest, doubtless did much to limit infl uences
unfavourable to theforest ecosystems state.
The 17th and 18th centuries
Palynological data make it clear that, in the17th–18th centuries, broadleaved (and
especially oak-lime-hornbeam) forests experienced apartial regression. Inthe pro-
fi les, adecline in theshare of tree pollen (mainly hornbeam and oak) is to be noted
(Figs. 3, 5and 6), albeit with Scots pine remaining at ahigh level (Fig.10), and with
theshare of spruce clearly on theincrease (Fig.9). Theshare of lime is relatively
large (Fig.7). Profi les also manifest aradical increase in thefrequency and taxo-
nomic diversity of anthropogenic indicators, with this suggesting that thechanges
in forest cover were rooted in human activity (Figs. 3, 4and 5). Ahigh frequency of
fi res is documented by theconsiderable presence of charcoal microparticles, as well
as thepollen of palynological indicators of habitat change due to theaction of fi re.
Inthe case of more nutrient-poor habitats in particular, this indicates anexpansion
of light-demanding plants, mainly heather (Fig.12). Another group of plant commu-
nities which expanded at this time was ruderal vegetation (distinct increase of
Arte-
misia
pollen; Fig. 12), with this pointing to increased synanthropisation of thelocal
fl ora.

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
The results of pollen analysis ilustrate thenatural effects of historically documented
forms of anthropopressure on thearea of theBiałowieża Forest (
i.a.
S
etal.
2013a). Historical data show that due to more liberal royal regulations and anincreas-
ing demand for forest products, theBiałowieża Forest of the17th and 18th centuries
(also thearea of thetoday’s Strict Reserve) experienced amarked increase in forest
exploitation. There was also adevelopment – more intensive than hitherto – of forest
beekeeping (H 1939; S 2006), getting bog iron ore (S 2009),
potash- and charcoal-burning, as well as tar-making (S
etal.
2013a). Traces of
this small-scale industrial activity are legible in theforest landscape of today (K
2007; S
etal.
2013a), with pollen sites BIA/314D and BIA/369F (Figs. 3and 4)
being located in theimmediate vicinity of where this activity was concentrated.
In connection with the demand for labour, new settlements mainly grew up on
themargins of theForest, though there were also hamlets in today’s Strict Reserve
area as such (S
etal.
2013b). Theexistence of apresumed settlement of pot-
ash-burners close to site BIA/314D is indicated by theovergrown remnants of fi eld fur-
rows mentioned by F (1986), among which fragments of modern-era ceramics
have been found. Another settlement was in existence in thevicinity of site BIA/369F.
In this area, known as
Uroczysko Paharelec
, it is possible to see the remnants of
aseries of hollows in theearth containing fragments of charcoal, as well as traces of
former tree cutting in thesurrounding forest. References to thelocality suggest that
there was asmall village of forest craftsmen here in the18th century (K 2007;
B 2015). Lithological composition and theresults of pollen analysis for theprofi le
of site BIA/369F (Fig.4) combined with radiocarbon dating indicate that deposition of
organic sediments began here in the16th–17th centuries, directly on apure sand lining
thefl oor of ashallow (50cm) depression. This, most probably, artifi cial form might
came forth due to diging up sand to reinforce anearby road. Inthis profi le, asharp limit
between mineral and organic sediments is characterised by thepresence of many
charcoal particles, as well as numerous spores of thebryophyte
Anthoceros puncta-
tus
– aspecies typical for communities of small therophytes spreading on moist sandy
substrata, and classifi ed to theIsöeto-Nanojuncetea class (P 1999). Asubse-
quent rise in thewater table caused ashallow body of water to come into existence
here, with this then becoming overgrown by wetland vegetation. High frequencies of
coprophilous fungi remains (especially
Cercophora
and
Sordaria
) suggest that thesite
might have been awatering area for livestock. Theresults of thepollen analysis from
thesite BIA/369F profi le point to alow share of hornbeam in theforest during 17th–18th
centuries, with this presumably denoting theintensive use of wood from this species
in theproduction of charcoal and potash (cf. S
etal.
2013a). Theproportions
of oak pollen are rather high, it is also worthy to note aquite marked and balanced

 M. Zimny, M. Latałowa, A. Pędziszewska
spruce contribution and relatively high frequency of lime pollen. These data suggest
selective harvesting of timber. Disturbances in thelocal environment are illustrated by
theregular occurrence of forest heliophyte pollen and charcoal microparticles.
The development of settlement forced anincrease in local food production, but still
relatively low (considering this period) proportions of cereal, buckwheat and segetal
weed pollen show alimited scale of cultivation in thestudy area. Theshare of pol-
len contributed by plants typical of open meadows and pastures in mesic habitats is
rather modest as well. Onthe other hand, acharacteristic feature of thepollen dia-
grams representing this period is therelatively large proportion of meadowsweet (
Fil-
ipendula
) pollen, which points to thepresence of wet meadows whose development
might in some part have been aconsequence of theexploitation of bog iron followed
by mowing. Thehigh share of pollen of heather and other forest heliophytes presum-
ably illustrates, not only habitat change due to theprocessing of forest products, but
also deliberate burning of undergrowth to support grazing (S and J
2004; B 2012). Thehigh incidence of fi res in theperiod under discussion has also
been documented by adendrochronological study (N
etal.
2010).
The 19th century
All of thepollen profi les register themarked change taking place in theforests of what
is today theStrict Reserve and its immediate vicinity at thebeginning of the19th century.
Across pollen spectra, thecontribution of hornbeam increases, with anupward trend also
noted for oak in most profi les. Incontrast, theproportions of lime pollen had tended to
go down. Ahigh level of occurrence of spruce pollen is maintained (orelse levels even
increase in some profi les); there is alack of palynological evidence for any marked recov-
ery of elm, ash or hazel populations. Proportions of Scots pine pollen and those of micro-
charcoal decline. Proportions of non-arboreal pollen are maintained at thesame level as
previously, or are even somewhat higher (Fig.10), but thestructure of this group of taxa
distinctly changes. First and foremost there is adecline in theproportions of
Calluna
nad
Artemisia
(Fig.12) concurrent with anincrease in thepollen of cultivated plants (Fig.11)
and of sheep’s sorrel (
Rumex acetosella
-t.); thelatter presumably spread in rye fi elds.
The palynological data offer aperfect refl ection of thefacts recorded in historical doc-
uments for this period, which describe negative impacts of theeconomic utilisation of
Białowieża Forest resources in the17th–18th centuries and subsequent restrictions intro-
duced in thelate-18th and early-19th century, including aprohibition on all forms of eco-
nomic activity associated with arisk of fi re (S 2006). Areduction in thefrequency

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
of fi res post-1781is made clear by thedendrochronologial data as well (N 2010).
Historical sources show that, as aresult, by the1920s, Scots pine had made aclear
retreat from thestands of theBiałowieża Forest, while therole of spruce – and subse-
quently of broadleaved trees – had increased (S 2006; N
etal.
2010).
Thesources in question describe theprocess by which forests regenerated, with these
being very visible in pollen diagrams (Fig.5). Historical documents also show how new,
small settlements with agricultural hinterland came into existence in the19th century
(F 1986; M
etal.
2013). This offers some explanation for theapparent
paradox present in thepollen diagrams, wherein apremise pointing to theclear regen-
eration of forest cover exists side by side with others showing how agricultural activity
increased (cf.Fig. 11).
The 20th century
The most recent history of theforests in theStrict Reserve of Białowieża Forest – about
thelast 100years – is well-known, thanks to theresults of ecological research, as well as
arich literature concerning their utilisation and protection. Acompendium of knowledge
on this subject is provided by theextensive monograph authored by J.F (1986)
that is concerned with theentire area of theForest with particular account of fi ndings
from theStrict Reserve. Beyond that, recent years have brought anincreasing number
of new and interesting publications concerned with thedynamics of forest communities
(
i.a.
K
etal.
2010; B
etal.
2011; B 2012). Some of these studies are based
on observations made on permanent plots, of which some were established more than
80years ago (
i.a.
B
etal.
1998; K 2005, 2007; B 2008; C
2008).
Where theperiod in question is concerned, thepollen diagrams provide only supple-
mentary data to theknowledge based on studies on recently ongoing processes. Onthe
other hand they enable direct comparisons of thepalynological record with documented
natural and historical facts. Themost characteristic feature of theyoungest parts of
the pollen profi les is a decline in anthropogenic indicators. Thelimited frequency of
cereal pollen (Fig.11) and that of fi eld weeds (
i.a.

R. acetosella
-t.) illustrates adecline in
area of cultivated fi elds in thevicinity of theReserve. There is adrop in thepercentage
values of palynological indicators of grazing (
i.a.

P. lanceolata
) (Fig.5), what conforms
with thedata elaborated by S
etal.
(2016), who assert that in theperiod from
theend of the19th century through to themid-20th century grazing was confi ned to just
asmall area along thesouthern boundary of today’s Strict Reserve. According to thehis-
torical data theintensity of this type of forest use in thesurroundings of theReserve

 M. Zimny, M. Latałowa, A. Pędziszewska
was low over thewhole period of thelast 100years. Thecurbing of agricultural activity
in thearea adjacent to today’s National Park was caused, not only by economic changes,
but also by successive demographic crises due to both wars and political turmoil of other
kinds (F 1986; J
etal.
1997; B 2001).
The 20th century was above all atime of the functioning of the forest communities
under conditions of legal protection. Reserve protection was ushered in here in 1921,
with thescope increasing up to 1929, to gradually take in thearea in thefork between
theRivers Narewka and Hwoźna. In1932, this area took on thestatus of the“National
Park at Białowieża”, only to become Białowieża (Białowieski) National Park by virtue of
adecision taken in 1947 (O 2002). This allowed for thespontaneous regeneration
of forest communities in places where they had beeen used previously. These ecological
processes took place in condition entirely unencumbered by direct human infl uence.
In thelate 19th and early 20th centuries, one of thekey factors exerting aninfl uence in
transforming broadleaved forest communities developing on mesotrophic and eutrophic
habitats was ahuge increase in numbers of game (in the years 1892–1915in particu-
lar). This was caused by animal breeding and rearing carried out by theservices of
theTsar with aview to hunting being facilitated (J
etal.
1997). Excessive
numbers of ungulates led to degeneration in communities of broadleaved forest due to
destruction of undergrowth and consequent (atleast transient) curtailment of renewal
among broadleaved trees. This allowed for afurther stage in theprocess of popula-
tion expansion of spruce which is atree species avoided by browsing animals (F
1986). Thedominance of spruce, especially in oak-lime-hornbeam forest habitats, was
of atransient nature, however. Several decades on, amass dieoff of this species began,
with this accelerating theprocess by which broadleaved stands regenerated. Traces of
this process are still visible in forest communities of theStrict Reserve, in theform of
generational gaps among trees of certain species, as well as ongoing loss from thestand
of Norway spruce (F 1986; K 2007; B
etal.
2011). Theinterpretation
of these phenomena in thecontext of long-term successional change is thesubject of
heated dispute at present (
i.a.
B
etal.
2016; J
etal.
2016).
The beginning of the20th century also marked theonset of theindustrial-scale exploita-
tion of timber in theBiałowieża Forest, which
inter alia
took in part of today’s Strict
Reserve, wherein thestand species composition continues to display traces of felling
carried out in three parallel strips ca. 1km long and 100mwide (F 1986, 1988).
Onthe area that forms theReserve, this was abrief episode; theongoing process of
secondary succession results in agradual blurring of boundaries between theold felling
site and its surroundings.

The Late-Holocene history of forests in the Strict Reserve of Białowieża National Park
The courses of thepollen curves for trees in thediagrams presented here make ref-
erence to data also arising out of direct observation of processes ongoing in forest
communities, with these pointing to theexpulsion of spruce from oak-lime-hornbeam
forest habitat (F 1986). Inthe uppermost parts of theprofi les agreater share is
accounted for by hornbeam and oak, and to some extent also hazel. Thepollen curves of
spruce are unstable, with culminations followed by deep depressions coming together
as arather weak downward trend. Thefl uctuations can be assumed to refl ect – at least
in part – repeated 20th-century outbreaks of thespruce bark beetle (K 2002),
which in particular appears when droughts become prolonged and are thus able to
exert anegative impact on thecondition of spruce trees. Thehydrological droughts
returning in recent decades (P
etal.
2002) are also amain cause of thefall in
thewater table that has affected moist and swampy habitats, then having aknock-on
effect on thecomposition and structure of stands, not only in theStrict Reserve, but
also in remaining parts of theForest. Encroachment of trees on open peatland areas has
also taken place as aresult (K 2005; S and C 2005). Thepal-
ynological data illustrate this process effectively, with birch and pine seen to expand at
thesites studied (Fig.3and 5), at thesame time as alder in theforest communities of
theBiałowieża Forest is subject to adownward trend (Fig.9).
Summary
The results of palynological research presented here offer aninsight into the long-
term dynamics of forest communities and themost important forest-forming species
in thearea of theBiałowieża National Park’s Strict Reserve, over theperiod of thelast
2000years. During this time, theForest has developed under conditions in which climate
change occurred many times, and with levels of anthropogenic pressure also varying.
Themost intensive human activity in thearea of Strict Reserve took place in theRoman
Iron Age, as theWielbark Culture tribes settled here, and then again in the17th and
18th centuries, when aseries of forest industries were permitted. Though material for
thestudy was mostly sampled at sites in close proximity to many archaeological traces
of past human activity, the pollen diagrams indicate only a limited infl uence of that
activity on thenatural environment. Theregeneration of forest communities took place
here with no more major involvement of pioneer species, and thus led to areturn to
thesituation prior to disturbances, with this attesting to maintained continuity of for-
est habitats. What is noteworthy are therelatively small proportions (ifwe consider
theperiod) of indicators of cultivation. This is again suggesting local farming on just
avery limited scale what is in accordance with historical documents reporting on about
400-years restrictions imposed on settlement development in Białowieża Forest. This

 M. Zimny, M. Latałowa, A. Pędziszewska
was presumably themain factor allowing ecosystems in theForest to remain in acondi-
tion allowing them to be identifi ed as “natural” in line with theclassifi cation of theinter-
national group of experts dealing with forest terminology (B 2005).
Acknowledgments
The work was made possible by funding earmarked for the MNiSW/NCN (N N305
167839) project. The opportunity to carry out fi eldwork within BNP was thanks to
thesupportive view of our research plans adopted by theScientifi c Council and Direc-
tor’s Offi ce of Białowieża National Park.
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304 B. Brzeziecki, A. Keczyński, J.Zajączkowski
Kamil Bielak, PhD, Eng.
Department of Forest Silviculture,
Warsaw University of Life Sciences
Leszek Bolibok, Associate Professor
Department of Forest Silviculture,
Warsaw University of Life Sciences
Bogdan Brzeziecki, Professor
Department of Forest Silviculture,
Warsaw University of Life Sciences
Włodzimierz Buraczyk,
Associate Professor
Department of Forest Silviculture,
Warsaw University of Life Sciences
Stanisław Drozdowski,
Associate Professor
Department of Forest Silviculture,
Warsaw University of Life Sciences
Leszek Gawron, PhD, Eng.
Department of Forest Silviculture,
Warsaw University of Life Sciences
Andrzej Grzywacz,
Retired Professor
Department of Forest Protection
andEcology, Warsaw University
of Life Sciences
Andrzej Keczyński, MSc, Eng.
Białowieża National Park
Anna Pędziszewska, PhD
Laboratory of Palaeoecology
andArchaeobotany, Department
of Plant Ecology, University of Gdańsk
Małgorzata Latałowa, Professor
Laboratory of Palaeoecology and
Archaeobotany, Department
of Plant Ecology, University of Gdańsk
Szeligowski Henryk,
Associate Professor
Department of Forest Silviculture,
Warsaw University of Life Sciences
Andrzej Szczepkowski,
Associate Professor
Department of Forest Protection
and Ecology,Warsaw University
of Life Sciences
Jacek Zajączkowski, PhD, Eng.
Department of Forest Silviculture,
Warsaw University of Life Sciences
Marcelina Zimny, PhD
Laboratory of Palaeoecology
andArchaeobotany, Department
of Plant Ecology, University of Gdańsk
Authors