Continuity and change in the vegetation of a Central European oakwood

Abstract

The issue of continuity in deciduous oakwood vegetation has been in the forefront of woodland ecological studies for many decades. The two basic questions that emerge from existing research are whether or not oakwoods can be characterized by long-term stability and what may be the driving forces of the observed stability or change. To answer these questions in a well-defined case study, we examined the history of a large subcontinental oakwood (Dúbrava) in the southeastern Czech Republic with interdisciplinary methods using palaeoecological and archival sources. Palaeoecology allowed us to reconstruct the vegetation composition and fire disturbances in Dúbrava in the past 2000 years, while written sources provided information about tree composition and management from the 14th century onwards. The pollen profiles show that the present oakwood was established in the mid-14th century with an abrupt change from shrubby, hazel-dominated vegetation to oak forest. This change was most probably caused by a ban on oak felling in AD 1350. From the 14th to the late 18th centuries Dúbrava had multiple uses, of which wood-pasture and hay-cutting kept the forest considerably open. The second remarkable change was dated to the late 18th century, when multiple-use management was abandoned and Dúbrava was divided into pasture-only and coppice-only parts. The last major shift occurred in the mid-19th century, when modern forestry and Scotch pine plantation became dominant. We conclude that Dúbrava Wood did not show stability in the long run and that its species composition has dramatically changed during the last two millennia. The most important driving force in the shaping and maintenance of the unique vegetation of Dúbrava was human management.

Full text

The Holocene
23(1) 46 –56
© The Author(s) 2012
Reprints and permission:
sagepub.co.uk/journalsPermissions.nav
DOI: 10.1177/0959683612450200
hol.sagepub.com
Introduction
Deciduous oakwoods have been on the decline for a century all
over Europe, because oaks fail to regenerate. This phenomenon is
often referred to as ‘Oak Change’ or ‘Oak Decline’ (Rackham,
2008; Watt, 1919). Available studies attribute it to various plant
diseases (e.g. Microsphaera alphitoides, Phytophthora species),
decline in human management or a combination of various fac-
tors (Jung et al., 2000; Luisi et al., 1993). Nonetheless, the term
‘oakwood’ denotes more than the dominance of oak species in the
tree layer – deciduous oakwoods are specific biotic assemblages
with many species more or less confined to this type of biotope
(Ellenberg, 1996; Konvička et al., 2004).
The issue of continuity in deciduous oakwood vegetation has
been in the forefront of woodland ecological studies for many
decades. The two basic questions that emerge from existing
research are whether or not oakwoods can be characterized by
long-term stability and what may be the driving forces of the
observed stability or change. Some scientists argued that decidu-
ous oakwoods are the natural vegetation of the relatively warm
and dry areas of Europe (Bohn and Neuhäusl, 2000; Ellenberg,
1996; Zólyomi, 1957), and are therefore stable communities
occupying a restricted range of ecological conditions for several
millennia. Others claimed that human management (burning, pas-
turing, litter raking and coppicing) rather than natural conditions
played a decisive role in the maintenance of European deciduous
oakwoods. This view is supported by the recent notable decline of
oakwood species assemblages and the ‘invasion’ of hornbeam,
maples, ash and lime following the abandonment of traditional
management (Hédl et al., 2010; Kwiatkowska et al., 1997;
Roleček, 2007). This opinion connects the stability of oakwoods
to continuity of management on a millennial case, which implies
fluctuations in vegetation with changes in human population den-
sities around particular oak forests.
While the question of stability versus change can be
approached using a single disciplinary perspective (such as paly-
nology, Huntley and Webb, 1988; Segerström, 1997; Ritchie,
1995), the study of driving forces requires a multiproxy approach
(Bürgi et al., 2004; Ireland et al., 2011; Pechony and Shindell,
2010). The comparison of natural scientific data with historical
written sources is particularly useful in this respect, since histori-
cal sources can provide a framework of interpretation for palaeo-
ecological data (Lindbladh et al., 2007; Veski et al., 2005).
Archival data have been successfully used to detect driving forces
of vegetation stability and change (Bürgi, 1999; Szabó, 2010b).
450200HOL23110.1177/095968361
2450200Jamrichová et al.The Holocene
2012
1 Institute of Botany of the Academy of the Sciences of the Czech
Republic, Czech Republic
2Charles University in Prague, Czech Republic
3Masaryk University, Czech Republic
Corresponding author:
Eva Jamrichová, Department of Vegetation Ecology, Institute of Botany
of the Academy of the Sciences of the Czech Republic, Lidická 25/27,
CZ-602 00 Brno, Czech Republic.
Email: eva.jamriska@gmail.com
Continuity and change in the vegetation
of a Central European oakwood
Eva Jamrichová,1,2 Péter Szabó,1 Radim Hédl,1 Petr Kuneš,2
Prˇemysl Bobek2 and Barbora Pelánková1,3
Abstract
The issue of continuity in deciduous oakwood vegetation has been in the forefront of woodland ecological studies for many decades. The two basic
questions that emerge from existing research are whether or not oakwoods can be characterized by long-term stability and what may be the driving
forces of the observed stability or change. To answer these questions in a well-defined case study, we examined the history of a large subcontinental
oakwood (Dúbrava) in the southeastern Czech Republic with interdisciplinary methods using palaeoecological and archival sources. Palaeoecology
allowed us to reconstruct the vegetation composition and fire disturbances in Dúbrava in the past 2000 years, while written sources provided information
about tree composition and management from the 14th century onwards. The pollen profiles show that the present oakwood was established in the
mid-14th century with an abrupt change from shrubby, hazel-dominated vegetation to oak forest. This change was most probably caused by a ban on oak
felling in ad 1350. From the 14th to the late 18th centuries Dúbrava had multiple uses, of which wood-pasture and hay-cutting kept the forest considerably
open. The second remarkable change was dated to the late 18th century, when multiple-use management was abandoned and Dúbrava was divided into
pasture-only and coppice-only parts. The last major shift occurred in the mid-19th century, when modern forestry and Scotch pine plantation became
dominant. We conclude that Dúbrava Wood did not show stability in the long run and that its species composition has dramatically changed during the
last two millennia. The most important driving force in the shaping and maintenance of the unique vegetation of Dúbrava was human management.
Keywords
ecosystem stability, historical ecology, management history, palynology, Quercus, temperate oakwood, written sources
Received 1 December 2011; manuscript accepted revised 30 April 2012
Research paper

Jamrichová et al. 47
In this paper we present the results of interdisciplinary research
on the long-term dynamics of a large lowland oakwood. This
wood is among the best preserved subcontinental deciduous oak-
woods in Central Europe (Roleček, 2007). Our study covers the
past two millennia, and we integrate (1) pollen and macrocharcoal
data from two small forest hollows, and (2) archival written docu-
ments on species composition and management at the site. We
aim to answer the following two questions:
(1) Is the oakwood at the study site characterized by long-
term stability in species composition or were there sig-
nificant changes during the past two millennia?
(2) What were the driving forces of stability or change?
Was the role of natural factors or human management
apparently more important?
Materials and methods
Study site
Hodonínská Dúbrava is a large (c. 3300 ha) and relatively well-
preserved ancient woodland (Figure 1). It is the core site of a spe-
cific type of subcontinental oakwoods, Carici fritschii-Quercetum
roboris (Chytrý, 1997; Roleček, 2007) – a community endemic to
this and a few adjacent sites. Because of low groundwater levels,
other vegetation types, such as alluvial forests (Alnion incanae)
and alder carrs (Alnion glutinosae) also occur. At higher and drier
elevations mesic oak-hornbeam woods (Carpinion) prevail.
Approximately half of the forest was turned into Scotch pine
plantations in the 19th and 20th centuries.
Dúbrava is located at the NW edge of the Pannonian Basin
(17°05΄00΄΄E, 48°52΄40΄΄N; Figure 1). Climate is ‘humid conti-
nental’ (cf. Peel et al., 2007). It is relatively warm and dry with c.
9°C of average annual temperature and 500–550 mm of precipita-
tion (Tolasz et al., 2007). The site is gently sloping towards the
SW, with elevations from 164 to 242 m. Dúbrava lies on the
fringe of the wide alluvium of the Morava river, on 150–200 cm
deep Quaternary blown sand deposits (Novák and Pelíšek, 1943).
Soils are extremely nutrient-poor arenic dystric cambisols (AOPK
CR 2002–2010), slightly acidic and prone to desiccation (Novák
and Pelíšek, 1943). The water-table fluctuates during the year.
The settlement history of the area is typical of Central European
lowlands. People have been living in the region since the Palaeo-
lithic with population peaks in the early Copper (4000–3400 bc),
late Bronze (1200–750 bc) and late Iron Ages (400–1 bc)
Figure 1. Location of Dúbrava Wood in the Czech Republic with the positions of previously published pollen profiles in the study area.

48 The Holocene 23(1)
(Měřínský and Šmerda, 2008a). Dúbrava Wood historically
belonged to the estate of Hodonín.
Palaeoecological analyses
The SW part of Dúbrava with numerous spring-submerged
depressions was surveyed in 2008. Guidance by R. Řepka helped
us locate 15 sites. We chose the six best-preserved sites for coring
using a Russian corer in 2008. All profiles were re-sampled for
pollen analysis in July 2009, mostly taken from a trench using 10
cm × 10 cm × 50 cm metal boxes. All profiles were analysed for
pollen at 5 cm resolution. Following the results of this prelimi-
nary analysis and radiocarbon dating (Table 1), we chose two pol-
len profiles denoted MS3 and MS4. The two sites represent
treeless wetlands of c. 50–100 m in diameter. Vegetation is domi-
nated by reed (Phragmites communis) at MS3 and by sedges
(Carex spp.) at MS4. The nearest vicinity of both wetlands is for-
est of Quercus petraea, Quercus robur and Pinus sylvestris.
Organic sediments from MS3 and MS4 stored in metal boxes
were subsampled by 1 cm. The preparation of samples for pollen
analysis followed standard techniques (Faegri and Iversen, 1989).
Samples containing mineral material were pre-treated with cold
concentrated HF for 24 h and then processed by KOH and acetol-
ysis. At least 500 pollen grains were identified using standard key
and photo collections (Beug, 2004; Reille, 1992, 1995, 1998); for
the determination of non-pollen palynomorphs we used van Geel
et al. (1980/1981). The nomenclature of pollen types follows
Beug (2004).
Macrocharcoal analysis was carried out on the same sections
as the pollen analyses. Sediments were sampled continuously at 1
cm increments using a calibrated sampler of 2 ml volume. The
macrocharcoal analysis of MS3 took place after the removal of
parts of the sediment for radiocarbon dating, therefore charcoal
data from some parts are missing. Extracted material was defloc-
culated with 10% KOH and subsequently non-charred organic
particles were bleached by 3% hydrogen peroxide (Schlachter
and Horn, 2010). Particles larger than 125 μm were separated
from the samples by wet sieving. Quantification of charred parti-
cles was performed using optical analysis of microphotographs
processed by ImageJ software (Rasband, 1997). Charcoal frag-
ments were identified according to their black colour and charac-
teristic shape (Enache and Cumming, 2006).
Pollen data are presented as percentages based on terrestrial
pollen sum, from which aquatics and local mire plants, pteri-
dophyta, algae, fungi and other non-pollen palynomorphs were
excluded. Charcoal concentration is expressed as the number of
pieces per 2 ml. Percentage pollen diagrams with macrocharcoal
histograms were created in Tilia v. 1.7.16 (Grimm, 2011). The
pollen profiles were divided into three (MS3) or four (MS4) pol-
len zones based on the results of ConsLink and visual analyses.
For a comparison of archival and pollen data, it is necessary to
understand the relevant source area of pollen (RSAP) (Sugita,
1994). In the case of small basins situated within forests Sugita
(1994) estimated the RSAP to be 50–100 m and about 40–50% of
pollen coming from trees growing within this radius, while the
rest (background pollen) originating from outside the RSAP. Cal-
cote (1995) confirmed this estimation by empirical research of
fossil pollen in forest hollows.
Selected plant macrofossils or charcoal taken from three
depths in each profile were used for AMS radiocarbon dating in
the Center for Applied Isotopes Studies, University of Georgia,
Athens GA, USA. An age–depth relationship model was con-
structed for both profiles using Clam v.1.0 R code (Blaauw, 2010;
R development Core Team, 2008). IntCal04 calibration curve
(Reimer et al., 2004) was used to calibrate radiocarbon dates. To
estimate the age of every pollen sample, linear interpolation
between the midpoints of calibrated dated levels was applied
(Table 1). The age–depth model was forced to go through 0 cm =
2009 bc/ad (bc ages are negative) (Figure 2). In this paper we use
calibrated years. The radiocarbon date of the organic sediment
from the base of the MS4 profile (5413 bc obtained from char-
coal) gave a result apparently too old when compared with the
MS3 profile, and was not used. This date may have resulted from
the sandy base allowing the vertical migration of plant macrore-
mains and charcoal. However, the date could possibly be correct,
which would refer to slow accumulation of organic material.
Nonetheless, the continuous presence of Juglans, which is known
to be a Roman import in the region (Hajnalová, 2001), in both
profiles appears to indicate that the base of MS4 is unlikely to be
older than the Roman Period.
Archival research
To find out about the history of management and vegetation in
Dúbrava, we studied the written records produced by the Hodonín
estate administration from the 14th to the 20th centuries. All
sources are presently kept at the Moravian Archives (MZA) in
Brno. Written sources are precisely dated, which allowed us to
establish a firm chronology of management changes. When com-
pared with data on management, for most of the study period
information on vegetation was patchy and rather vague. However,
for the last two centuries precise data are available on tree species
composition. The following kinds of sources were used:
(1) Charters are records of legal transactions. The first
charters dealing directly with Dúbrava are from the
mid-14th century, while the last ones are from the 18th
century.
(2) Urbaria are conscriptions of all incomes from an estate.
Four urbaria survive that provide relevant information
about the Hodonín estate. They date from 1600 (MZA
F5 kniha 1a, analyzed in Chocholáč, 1994), c. 1654
(incomplete: MZA F5 kniha 1), 1691 (MZA F5 kniha 3)
and 1805 (MZA F5 kniha 4).
Table 1. Calibration of radiocarbon dates of the MS3 and MS4 profiles.
Depth (cm) Lab. code Sample ID Dated material 14C age (BP) Calibrated 14C age
Mean Range
40–38 UG-7745 MS3,1 Charcoal 2000 ± 25 ad 6 47 bc– ad 60
34–32 UG-7744 MS3,2 Carex seeds 610 ± 25 ad 1349 ad 1297–1401
14–12 UG-7743 MS3,3 Carex seeds 170 ± 20 ad 1808 ad 1665–1950
44–42 UG-8518 MS4 Charcoal 6420 ± 25 5413 bc 5472–5341 bc
35–33 UG-747 MS4,2 Seeds, charcoal 560 ± 25 ad 1368 ad 1313–1424
17–15 UG-7746 MS4,3 Carex seeds 120 ± 25 ad 1809 ad 1681 –1937
ad: Anno Domini; bc: Before Christ; BP: Before Present (1950).

Jamrichová et al. 49
Figure 2. Age–depth relationship of (a) MS 3 and (b) MS4 profiles based on three radiocarbon dates for each profile. Cal.: calibrated ages;
ad: Anno Domini (after Christ); bc: Before Christ.
(3) Estate conscriptions describe the value of landed prop-
erty in a given historical moment. A detailed conscrip-
tion survives from 1692 (MZA F5 kniha 5).
(4) Account books were kept by woodland owners to reg-
ister incomes generated by the cutting and selling of
underwood and timber. For Dúbrava only a few remain,
covering the years 1765–1772 (MZA F5 karton 538).
(5) Forestry management plans (FMP) are detailed sur-
veys containing information on the name, size and
position of each woodlot, as well as on tree composi-
tion and forest structure, supplemented by current and
planned management. FMPs of Dúbrava are from 1851
(MZA F5 kniha 232), 1864 (MZA F5 kniha 233-236,
238, 240) 1906 (MZA F5 kniha 242, 244, 245), 1925
(MZA F263 kniha 1), 1936 (MZA F263 kniha 4, 5) and
1952 (MZA F302 kniha 4).
(6) Forestry documents were produced by the local for-
estry administration. Such documents include e.g. vari-
ous surveys of woodland areas, discussion on types of
management and detailed diaries of yearly activities.
(7) Large-scale maps (c. 1: 30,000 and larger) are gener-
ally available in the Czech Lands from the 18th century
onwards. We used three nationwide surveys prepared
by the Austrian Army in 1764–1783, 1836–1852 (both
1: 28,800) and 1876–1880 (1: 25,000), available online
at http://oldmaps.geolab.cz. A much more detailed
(1: 2880) set of maps were drawn in 1824–1843 as part
of the so-called Stable Cadastre (Bičík et al., 2001).
Finally, maps produced as parts of forestry manage-
ment plans were used. Three sets of maps survive from
Hodonín: 1851 (MZA F5 mapa 54–59), 1884 (MZA F5
mapa 60–64, note that the FMP itself is lost) and 1906
(MZA F5 mapa 65–70).
Results
Pollen stratigraphy and vegetation history
Based on changes in dominant taxa (Table 2), the pollen diagrams
were divided into three (MS3) or four (MS4) phases. The first
three phases are highly similar in both profiles, while the fourth
one, covering the second half of the 20th century, is visible only
in MS4 (Figures 3 and 4). Historical documents show correspond-
ing results; major management changes happened approximately
in the transition periods between the pollen zones. Because of the
high similarity between the pollen spectra and historical sources,
the results are presented jointly.
Roman period to the high Middle Ages (1st century ad–mid-14th
century ad) – Phase 1. At the beginning of the sedimentation
process dated to the 1st century ad, Dúbrava consisted mainly of
light-demanding woody species, such as Corylus (19%) and Bet-
ula (17%). The relatively low proportion of tree pollen and of
Juniperus (3%) also suggests open forest vegetation. A high
amount of pollen of coniferous trees, such as Picea (18%) and
Abies (6%) was also recorded. Corylus produces little pollen as an
understorey species (Rackham, 1988). The observed high amount
of Corylus in the profile suggests that it had a dominant position
in the vegetation (Gardner, 2002), and that its pollen spread was
hardly hindered by overstorey vegetation. This and the presence
of Juniperus allow us to interpret the vegetation in the nearest
vicinity of the study site as shrubby woodland. Such vegetation is
implied in the original, high medieval name of Dúbrava, which
was Klečka. This name was first recorded in a charter interpola-
tion from ad 1350 (Boček, 1839: 204–205). In Old Czech,
‘klečka’ refers to a place with shrubs (Gebauer, 1970). The basal
layers of both profiles contained low amounts of macroscopic
charcoal particles. These provided no evidence for local fire
events which could have been connected to the shrubby vegeta-
tion dominated by Corylus (cf. Tinner et al., 2005).
The composition of herbaceous pollen in Phase 1 indicates the
presence of an intensively managed landscape. The high amount
of Artemisia, Asteraceae, Silenaceae, Rubiaceae and Valeriana
officinalis-type implies meadows. The constant occurrence of
Cerealia indicates the presence of arable fields in the nearest
vicinity of the site. Ruderal plants such as Polygonum aviculare-
type and Polygonum persicaria-type indicate trampled habitats
and cultivated land (Behre, 1981; Gaillard, 2007). At the end of
Phase 1, the pollen of Plantago lanceolata-type, Rumex acetosa-
type and Melampyrum appeared. These could be connected with
grazed grasslands.

50 The Holocene 23(1)
Table 2. Characteristics of main phases based on changes in pollen taxa and historical events.
Cal. ad MS3 MS4 Written sources
Max. % Min. % Max. % Min. %
Phase 4
1941
1808/1809
1349 /1348
6
n/a n/a Betula, Populus
Cerealia,
Zea mays,
Typha latifolia t.
Artemisia
Chenopodiaceae
Cyperaceae
Dominance of high forests, afforestation of open
areas, abandonment of drainage canals
End of multiple
management,
separation of
pasture and
woodland
Ban on cutting oaks
Phase 3
Loranthus europaeus
Poaceae, Equisetum
Corylus Picea
Artemisia, Asteraceae,
Scrophulariaceae, Rubiaceae,
Cyperaceae, Typha latifolia t.
Fraxinus
–
Corylus
Rubiaceae
Strict coppicing regime in woodland later to be
replaced by high-forests, grubbing out of one third of
the Wood and turning it into pasture. Digging of of
drainage canals. Beginning of Pinus plantations.
Phase 2
Quercus
Betula, Pinus
Chenopodiaceae, Asteraceae,
Rubiaceae, Rumex acetosa t.,
Rhinanthus,
Polygonum persicaria t.,
Cerealia, Secale,
Podospora, Sporormiella
Tilia, Alnus
–
Quercus
Pinus
Chenopodiaceae, Asteraceae,
Brassicaceae, Lotus t., Rubiaceae ,
Filipendula,
Chaerophyllum hirs.
Plantago lanceolata t.,
Picea, Ulmus, Alnus
–
Multiple-use management: wood-pasture, hay
meadows, coppicing, pannage, wild-fruit and oak gall
collection, beehives and arable fields in the Wood.
Gradual change in name from Klecˇka (‘place with
shrubs’) to Dúbrava (‘oakwood‘)
Polygonum aviculare t., Secale
Phase 1
Corylus, Picea, Abies, Ulmus,
Tilia, Juniperus,
Alnus
Artemisia, Silenaceae, Senecio
t., Scrophulariaceae,
Valeriana off. t.,
Polygonum aviculare t.,
Polygonum persicaria t.,
Cyperaceae,
Typha latifolia t., Sparganium t.
Quercus, Pinus
Cerealia, Secale
Corylus, Picea, Tilia, Ulmus,
Juniperus, Alnus
Silenaceae, Ranunculaceae,
Senecio t.,
Apiaceae
Geranium,
Polygonum persicaria t.
Quercus, Pinus
Cerealia, Secale Original name Klecˇka refers to shrubby vegetation
Cal.: calibrated ages; ad: Anno Domini (after Christ); Max%: maximum (or increasing) of pollen percentage; Min%: minimum (or decreasing) of pollen percentage.

Jamrichová et al. 51
Figure 4. Percentage pollen diagram of selected pollen types with macrocharcoal concentrations from MS4 core.
Figure 3. Percentage pollen diagram of selected pollen types with macro-charcoal concentrations from MS3 core. Parts of sediment without macrocharcoal analysis are marked as No Data.

52 The Holocene 23(1)
Late Middle Ages to the early Modern Period (mid-14th century
ad–end of 18th century ad) – Phase 2. The first great change in
forest composition occurred in the mid-14th century. It was
recorded in both profiles and is dated ad 1348 (MS3) and ad 1369
(MS4). This event is characterized by a rapid increase in Quercus
(30%, i.e. by 20%), and in Pinus (to 10%). It was accompanied by
a strong decline in Corylus (6%), Abies (1%) and Picea (4%). Bet-
ula did not change. The dominance of oak, birch and pine has been
characteristic for Dúbrava ever since the beginning of Phase 2.
Two charters refer to the active protection of oaks in Dúbrava
precisely in this period. The first one is the above-mentioned
interpolated charter from 1350, which gave the citizens of
Hodonín the right to take dry wood and grass in the Wood but
forbade them to fell living oaks (Boček, 1839: 204–205, for a
discussion on the dating of the charter, see Měřínský and Šmerda,
2008b). The other one is the foundation charter of the Augustin-
ian monastery in Brno from ad 1370. It was included in this
document that the tenants of the monastery had the right to cut
timber and firewood in Dúbrava ‘with the exception of oak trees,
which they must not cut down at all’ (translation from Latin orig-
inal, MZA F5 karton 11 inv. č. 744, fol. 25–32). The ban on cut-
ting oaks was included in a number of privileges in later centuries
as well (e.g. 1531 – MZA F5 karton 3 inv. č. 29; 1600 urbarium –
MZA F5 kniha 1a).
In Phase 2 the maxima of Quercus in both profiles (34% MS3;
28% MS4) and the highest percentage of AP (78%) were recorded.
The subsequent decrease in Quercus pollen is synchronous with
an increase in the pollen of the other main tree species: Carpinus,
Fagus, Abies and Picea. From the middle of the 14th century
onwards, the general increase in Cerealia, the start of a continu-
ous pollen curve of Secale and the occurrence of weeds (Centau-
rea cyanus – a typical high-medieval weed, Papaver rhoeas-type)
may be connected with the expansion of arable land as a conse-
quence of population growth in the surrounding area. The
unchanged curves of Plantago lanceolata-type, Rubiaceae, Sene-
cio-type, Rumex acetosa-type, Chenopodiaceae and Lotus-type
suggest the continuous presence of grasslands and pastures. This
argument is also supported by findings of spores of coprophilous
fungi (Podospora, Sporormiella) in the sediment. The pollen of
Juniperus and the spores of Sporormiella are indicators of live-
stock farming (Davis, 1987).
We observed a gradual increase in macroscopic charcoal par-
ticles throughout Phase 2. This could be connected with higher
fire susceptibility of the vegetation caused by the expansion of
Pinus or with the intensification of human activities in the vicinity
of the study sites. Higher concentrations of charcoal particles at
the end of this phase in both profiles and a slight decrease in
Quercus and increase in Betula and Cyperaceae may indicate a
distinct fire event, however, the percentages of Quercus, Betula
and Cyperaceae swiftly returned to their previous values.
In this period the name of the Wood changed. The original
Klečka (‘a place with shrubs’) was gradually replaced by Dúbrava
(‘oakwood’). ‘Dúbrava’ was first used as a common noun that
added information on the Wood (e.g. 1370: ‘the oakwood
[dúbrava] that is called Klečka’ – MZA F5 karton 11 inv. č. 744,
fol. 25–32), and only later became a geographical name. The last
occurrence of the name Klečka is from 1531 (MZA F5 karton 3
inv. č. 29), after that only Dúbrava was used. Two urbaria (1600 –
MZA F5 kniha 1a; 1691 – MZA F5 kniha 3) provide detailed
information on the management of Dúbrava. The most character-
istic feature was multiple use, which included wood-pasture, pan-
nage (the fattening of domestic pigs on acorns), hay cutting in
woodland meadows, firewood cutting and the collecting of straw-
berries and oak galls. There were managed ponds, beehives and
even arable fields within the Wood. The system was complex but
not random: every use was carefully regulated temporally and
spatially. The urbaria also include a description of the boundaries
of Dúbrava, from which it is clear that the boundaries of the Wood
were similar to those of today and that a few hundred metres from
the pollen sites we investigated there were arable fields. Rela-
tively little direct information is available on tree species in this
period: apart from frequent references to acorns and oak galls, the
1692 estate conscription (MZA F5 kniha 5) mentioned that
Dúbrava comprised mostly of oaks and partly of aspen and birch.
The lack of Pinus in this list is noteworthy.
Modern Period (19th century ad–present) – Phases 3 and 4. At
the beginning of the 19th century the second major change in the
composition of Dúbrava can be observed. In the pollen diagrams
this was reflected by a change in tree composition. A moderate
decline in Quercus and increase in Betula, Pinus, Salix, Populus
and Fraxinus can be associated with several, possibly interacting
factors. A significant decline in Cerealia and Secale, dated to the
beginning of the 19th century, is visible in MS3 but not in MS4;
the latter is characterized by an increase in Polygonum aviculare-
type, Trifolium repens-type and large quantities of coprophilous
fungal spores (Podospora). The apparent contradiction (increase
in human activities in MS4, decrease in MS3) can be explained by
the different location of the profiles within the Wood.
The last change was recognized only in MS4 and refers to
Phase 4. It is dated to the mid-20th century and is characterized
by a rapid increase in Populus and Betula and the disappear-
ance of pasturing indicators, such as Plantago lanceolata-type,
Trifolium repens-type and Rubiacae. It refers to the abandon-
ment of pastures and fields inside or immediately outside the
Wood and secondary succession with Populus and Betula as
pioneer trees. The curve of Quercus rose again but it never
reached the medieval maximum of the beginning of Phase 2. At
the end of this phase we also recorded a decrease in the curves
of other herbs, for example Filipendula, Potentilla-type, Chae-
rophyllum hirsutum, Silenaceae.
Macroscopic charcoal concentrations steeply increased towards
the top of MS4. This indicates the presence of fire events in the
fire-prone Pinus stands which were established by modern for-
estry. The increase in charcoal particles could also be associated
with the construction of a railway line along Dúbrava Wood. The
first steam engine on the Emperor Ferdinand Northern Railway
passed through the town Hodonín in ad 1841 (Vykoupil, 2008).
Written sources show a major management change at the
beginning of Phase 3. In the late 1780s Dúbrava was divided into
two parts. The larger part (c. two-thirds of the whole) was
enclosed by a woodbank (Szabó, 2010a) and turned into a cop-
pice, while on the remaining one-third all trees were quickly
removed and the area was turned into pasture. The former mul-
tiple management was abolished; pasturing, hay-cutting and pan-
nage were banned. By the mid-19th century, however, the
forestry administration changed its mind about management and
started transforming the coppices into high-forests, partly with
the help of Pinus plantations. Timber producing high-forests
(including plantations) have formed the majority management
type since the 1950s (as witnessed by consecutive forestry man-
agement plans: 1851 – MZA F5 kniha 232; 1864 – MZA F5
kniha 233-236, 238, 240; 1906 – MZA F5 kniha 242, 244, 245;
1925 – MZA F263 kniha 1; 1936: MZA F263 kniha 4, 5; 1952 –
MZA F302 kniha 4). There were continuous efforts to afforest
the open areas in the Dúbrava, however, these were unsuccessful
until the 1920s. The pastures created in the late 18th century did
not last long, and by 1906 (MZA F5 kniha 242, 244, 245) they
reverted back to woodland (Figure 5).
The hydrological regime of the Wood was also changed. Some
time in the 19th–20th centuries much of the territory of Dúbrava
was drained by a network of channels. These channels connected

Jamrichová et al. 53
wetter areas and drained their water into larger canals that had
been created in and around the Wood since the 15th century.
These drainage channels are still visible in the field but are not
maintained any more. Their dating is uncertain. The 1925 FMP
(MZA F263 kniha 1) mentioned maintenance work on canals that
were created before 1907, therefore we assume that the channels
were dug already in the 19th century.
Discussion
The combination of pollen analysis, macrocharcoal analysis and
the study of historical documents provides synergetic results about
vegetation stability and historical human impact in Dúbrava Wood.
Palaeoecological analyses enabled us to reconstruct vegetation
composition and fire disturbances during the past 2000 years while
written archival material revealed information on tree composition
and management practices for the past seven centuries.
Stability/change in species composition in a regional
context
Our results show that in the study period the species composition
of Dúbrava went through significant changes. In the 14th century
the vegetation of Dúbrava Wood changed from shrubby growth
composed mainly of Corylus and Betula to subcontinental oak-
wood. In the past two centuries mesophilous species started to
spread and Pinus plantations appeared – this process continues to
the present.
To gain a more general picture of forest vegetation develop-
ment in the study region, we used previously published palaeo-
ecological data from four nearby sites: Vracov (Rybníčková and
Rybníček, 1972; Svobodová, 1997), Svatobořice-Mistřín (Svo-
bodová, 1989, 1997), Anšův Dvůr (Svobodová, 1990) and Pohan-
sko (Doláková et al., 2010; Svobodová, 1990). They showed that
Quercus started to spread in the study region in c. 6900 bc. After
this date, grassy subxerophilous oakwoods or mixed oak-lime-
hornbeam forests developed in mesic conditions, and floodplain
forests prevailed in wet conditions. All four pollen profiles
recorded the dominant presence of Quercus from c. 6900 bc to the
early Middle Ages, c. 6th–9th centuries ad. Unlike at the other
three sites, there were relatively high amounts of Fagus and Abies
at Vracov from c. 3900 bc onwards. This may have resulted from
long-distance transport as the site represents a larger lake basin. In
contrast to the vegetation recorded in the four profiles, from the
1st to the 14th centuries ad Dúbrava consisted mainly of Corylus
and Betula. Quercus, Tilia, Ulmus and Carpinus occurred only as
admixture species. However, we cannot tell whether the vegeta-
tion observed at the beginning of the study period had been stable
in previous centuries or whether it was the result of recent
changes.
The most significant spread of oak in Dúbrava was recorded at
the beginning of the 14th century. This process was accompanied
by a decrease in other trees and shrubs, mainly Corylus. A com-
parison with other pollen diagrams from the region shows a simi-
lar decrease in Corylus and increase in Quercus, however, at a
completely different date, in c. 1200 bc (Svobodová, 1997). The
massive spread of oak recorded in Dúbrava has no analogues in
southern Moravian pollen profiles in this period, all four of which
show a decline in Quercus starting from the early Middle Ages.
The pollen profiles from Dúbrava ended with a visible decline
in oak and an increase in ash, birch and pine in the 19th century.
This increase in Fraxinus and Pinus was recorded in other pollen
diagrams from the study region as well. The spreading of Fraxi-
nus in predominantly oak forests is probably a natural reaction to
the absence of organic matter removal and is part of a gradual
change to a shady mesic forest (Hédl et al., 2010; Hofmeister et al.,
2004). Pinus spread mainly as a result of plantation forestry.
Driving forces of stability/change in species
composition
A possible explanation of the rapid spread of Quercus in Dúbrava
in the 14th century ad could be provided by the onset of the ‘Little
Ice Age’ (LIA), which began in c. ad 1300 (Matthews and Briffa,
2005). Climate change (cooler and moisture conditions) could
have influenced vegetation composition and cause the spreading
of Quercus, which is less sensitive to late frosts than Fagus. In
Białowieża forest (Poland), Mitchell and Cole (1998) attribute the
dominance of Quercus to a competitive edge in edaphic condi-
tions. However, Faliński (1986) described the dominance of
Quercus in this forest as a result of grazing (herbivores and cat-
tle). After a reduction in grazing, Quercus was replaced by Carpi-
nus. In our study region, no other pollen profile records an
increase in Quercus in the LIA; climate change is therefore
unlikely to have caused the dramatic change in the vegetation of
Dúbrava. Another possible explanation of the massive spread of
oak is fire. Recent studies form North America show that the
widespread occurrence and dominance of oak is the result of fre-
quent fires (Abrams, 1992). Today, oak is in decline because of
fire suppression by humans, which leads to a gradual replacement
of oak by shade-tolerant species (Dey, 2002; Little, 1974; Lorimer,
1993; Van Lear, 1991). However, the results of macrocharcoal
analyses from Dúbrava showed no major fire event parallel with
Figure 5. Changes in the amount of (a) broadleaved forest, conifer
forest and open areas, (b) high forest, coppice and open areas and in
Dúbrava since the end of the 18th century. Data prior to 1851 are
approximate.

54 The Holocene 23(1)
the sudden change in species composition at the beginning of
Phase 2. There is some indication of a possible fire at the end
of Phase 2, nevertheless even in this case higher concentrations of
charcoal particles were not connected to any lasting influence on
species composition. Fire is therefore unlikely to have been a
driving force of species composition changes.
In a regional context, the changes in the species composition
of Dúbrava Wood appear to be rather exceptional. In such cases,
site history is often the best explanatory factor (Ejarque et al.,
2009; Lindbladh et al., 2007; Veski et al., 2005). Two written
documents mentioned the protection of oaks in the mid-14th cen-
tury, which precisely coincides with the pollen data. However, the
interpretation of these charters needs careful attention. Such bans
were a commonplace in medieval charters and did not necessarily
have to have any concrete consequences. In Sweden in the 18th
and 19th centuries a similar ban is known to have caused a decline
in oak numbers, although the socioeconomic conditions here were
very different from late medieval Moravia (Eliasson and Nilsson,
2002). Nonetheless, in our case the existence of two charters and
a simultaneous increase in oak pollen can hardly be a coincidence.
These charters indicate that from the middle of the 14th century
oaks were in fact actively protected in Dúbrava, which led to
changes in the vegetation. According the Vera Hypothesis (Vera,
2000), an increase in wood-pasture (also recorded in the two char-
ters) could also have promoted oaks. Similar conclusions were
arrived at for an earlier period at Pohansko, where the spread of
Quercus accompanied by a slight decrease in Tilia, Fraxinus and
Ulmus in c. ad 600–700 was attributed mainly to grazing by pigs
(Kratochvíl, 1981; Svobodová, 1990;).
From the 14th to the late 18th centuries, Dúbrava had multiple
uses, some of which (pasturing and hay cutting) kept the Wood
relatively open. In this phase the maxima of Quercus in both pro-
files (34% MS3; 28% MS4) and the highest percentage of AP
(78%) were recorded. This is confirmed by the 1692 estate con-
scription, which claimed that the Wood comprised mostly of oaks.
The change in species composition (slight decline in Quercus and
spreading of Fraxinus, Betula, Populus and Pinus) at the end of
this period could have multiple reasons. One is a change in hydro-
logical conditions (the construction of drainage channels), another
could be windbreak as suggested by an increase in pioneer trees
(Populus and Betula). Written documents show that multiple-use
management ended exactly at this time, and the part where the
forest hollows are situated was turned into pure coppice and some
50 years later into high-forest. Therefore the change in species
composition can also be associated with a reduction of grazed
areas (disappearance of pasturing indicators) and the gradual con-
version to high-forest. A similar situation was observed in
Białowieża forest (Faliński, 1986; Mitchell and Cole, 1998),
where species of open habitats gradually declined and mesophytic
tree species (Carpinus) replaced oak. This has lead to the loss of
species assemblages typical for subcontinental oakwoods (Kwiat-
kowska et al., 1997).
Protecting the unique vegetation of subcontinental
oakwoods
Open canopy oakwoods currently host many endangered species
(Spitzer et al., 2008). To sustain their populations, these species
had to find suitable habitats throughout the Holocene. It is often
argued that open oakwoods with wood-pasture continuity must
have been present since prehistoric times (e.g. Vera, 2000; Vodka
et al., 2009). While this is certainly possible, other management
types and tree species compositions could have provided equally
suitable conditions. For example Milovice Wood, a large subcon-
tinental oakwood not far from Dúbrava, was managed as a cop-
pice-with-standards for at least 600 years (Szabó, 2010b). Only
when coppicing ceased did the subcontinental character of the
vegetation begin to rapidly fade (Hédl et al., 2010). Our results
indicate that open woodland could have consisted of various com-
munity types. In Dúbrava, open forest communities were appar-
ently present from the first millennium ad, while the oakwood
fully developed only in the 14th century. This suggests that even
from the point of view of species sensitive to canopy openness,
open oakwoods might not have been the only option for survival
in the Holocene.
Conclusions
We conclude that Dúbrava Wood did not show stability in the
long run and that its species composition has dramatically
changed during the last two millennia. From ad 0 to 1350 Dúbrava
was almost certainly not an oakwood. The origins of its present
species composition date back to the 14th century, when inten-
tional management caused a shift from shrubby, hazel-domi-
nated vegetation towards an oakwood. The most important
driving force in the shaping and maintenance of the unique vege-
tation of Dúbrava was human management. medieval manage-
ment promoted oaks, and the open oakwoods were further
maintained by multiple-use management in the early Modern
Period. They were subsequently drastically reduced in extent by
modern forestry plantations continuing to date. The species in the
herb layer could have been present in the millennia preceding the
past 2000 years; however, until the 14th century they had to sur-
vive in other types of vegetation than subcontinental oakwoods.
Acknowledgements
We would like to thank Radek Řepka for introducing us to the
sites which were sampled for pollen. We are grateful to Dušan
Lekeš for his help during fieldwork and to Zuzana Formánková
for laboratory work.
Funding
This study was supported by grant no. IAA600050812 ‘Lowland
woodland in the perspective of historical development’ from the
Grant Agency of the Academy of Sciences of the Czech Repub-
lic and as a long-term research development project no. RVO
67985939.
References
Abrams MD (1992) Fire and the development of oak forests. BioScience 42:
346–353.
Behre KI (1981) The interpretation of anthropogenic indicators in pollen dia-
grams. Pollen et Spores 23: 225–245.
Beug HJ (2004) Lietfaden der Pollen bestimmung für Mitteleuropa und
angrezende Gebiete. München: Verlag Dr Friedrich Pfeil.
Bičík I, Jeleček L and Štěpánek V (2001) Land-use changes and their social
driving forces in Czechia in the 19th and 20th centuries. Land Use Policy
18: 65–73.
Blaauw M (2010) Methods and code for ‘classical’ age modelling of radiocar-
bon sequences. Quaternary Geochronology 5: 512–518.
Boček A (ed.) (1839) Codex Diplomaticus et Epistolaris Moraviae tom. 2.
Olomouc.
Bohn U and Neuhäusl R (2000) Karte der natürlichen Vegetation Europas.
Maßstab 1:2.500.000. Teil 1: Erläuterungstext mit CD-ROM; Teil 2: Leg-
ende; Teil 3: Karten. Münster: Landwirtschaftsverlag.
Bürgi M (1999) A case study of forest changes in the Swiss lowlands. Land-
scape Ecology 14: 567–75.
Bürgi M, Hersperger AM and Schneeberger N (2004) Driving forces of land-
scape change – Current and new directions. Landscape Ecology 19:
857–868.
Calcote R (1995) Pollen source area and pollen productivity: Evidence from
forest hollows. Journal of Ecology 83: 591–602.
Chocholáč B (1994) Hodonínske panství počátkem 17. století. Časopis matice
moravské 113: 59– 70.
Chytrý M (1997) Thermophilous oak forests in the Czech Republic: Syntaxo-
nomical revision of the Quercetalia pubescenti-petraeae. Folia Geobo-
tanica 32: 221–258.

Jamrichová et al. 55
Davis OK (1987) Spores of dung fungus Sporormiella: Incerased abundances
in historic sediments and before Pleistocene megafaunal extinction. Qua-
ternary Research 28: 290–294.
Dey DC (2002) The ecological basis for oak silviculture in eastern North
America. In: McShea WJ and Healy WM (eds) Oak Forest Ecosystems
Ecology and Management for Wildlife. Baltimore MD: The John Hopkins
University Press, pp. 60–79.
Doláková N, Rozsková A and Přichystal A (2010) Palynology and natural envi-
ronments in the Pannonian to Holocene sediments of Early medieval cen-
tre Pohansko near Břeclav (Czech Republic). Journal of Archaeological
Science 37: 2538–2550.
Ejarque A, Juliá R, Riera S et al. (2009) Tracing the history of highland
human management in the eastern Pre-Pyrenees: An interdisciplinary
palaeoenvironmental study at the Pradell fen, Spain. The Holocene 19:
1241–1255
Eliasson P and Nilsson SG (2002) ‘You should hate young oaks and young
nobleman’ The environmental history of oaks in eighteenth- and nine-
teenth-century Sweden. Environmental History 7: 659–677.
Ellenberg H (1996) Vegetation Mitteleuropas mit den Alpen. 5th edition. Stutt-
gart: Ulmer.
Enache MD and Cumming BF (2006) Tracking recorded fires using charcoal
morphology from the sedimentary sequence of Prosser Lake, British
Columbia (Canada). Quaternary Research 65: 282–292.
Faegri K and Iversen J (1989) Textbook of Pollen Analysis. 4th Edition. Chich-
ester: John Wiley & Sons.
Faliński JB (1986) Vegetation dynamics in temperate lowland primeval for-
est. In: Faliński JB (ed.) Geobotany. Dordrecht: Dr W Junk Publishers,
pp. 39–111.
Gaillard MJ (2007) Detecting Human impact in the pollen record. In: Elias SA
(ed.) Encyclopedia of Quaternary Science. Elsevier, pp. 2570–2595.
Gardner AR (2002) Neolithic to Copper Age woodland impacts in northeast
Hungary? Evidence from the pollen and sediment chemistry records. The
Holocene 12: 541–553.
Gebauer J (1970) Slovník staročeský. Praha: Academia.
Grimm EC (2011) Tilia software v.1.7.16. Springfield IL: Illinois State
Museum.
Hajnalová E (2001) Ovocie a ovocinárstvo v archeobotanických nálezoch na
Slovensku. Nitra.
Hédl R, Kopecký M and Komárek J (2010) Half a century of succession in a
temperate oakwood: From species-rich community to mesic forest. Diver-
sity and Distributions 16: 267–276.
Hofmeister J, Mihaljevič M and Hošek J (2004) The spread of ash (Fraxinus
excelsior) in some European oak forests: An effect of nitrogen depo-
sition or succesional change? Forest Ecology and Management 203:
35–47.
Huntley B and Webb T III (eds) (1988) Vegetation History. Handbook of Veg-
etation Science, vol. 7. Dordrecht: Kluwer.
Ireland AW, Oswald WW and Foster DR (2011) An integrated reconstruction
of recent forest dynamics in a New England cultural landscape. Vegetation
History and Archaeobotany 20: 245–252.
Jung T, Blaschke H and Oßwald W (2000) Involvement of soilborne Phytoph-
thora species in Central European oak decline and the effect of site factors
on the disease. Plant Pathology 49: 706–718.
Konvička M, Čížek L and Beneš J (2004) Ohrožený hmyz nížinných lesů:
ochrana a management. Olomouc: Sagittaria.
Kratochvíl Z (1981) Tierknochenfunde aus grossmährischen Siedlung
Mikulčice, I. Das Hausschwein. Studie Archeologického Ústavu ČSAV
Brno 9. Praha: Academie.
Kwiatkowska AJ, Spalyk K, Michalak E et al. (1997) Influence of the size and
density of Carpinus betulus on the spatial distribution and rate of deletion
of forest-floor species in thermophilous oak forest. Plant Ecology 129:
1–10.
Lindbladh M, Brunet J, Hannon G et al. (2007) Forest history as a basis for
ecosystem restoration – A multi-disciplinary case-study in a south Swed-
ish temperate landscape. Restoration Ecology 15: 284–295.
Little S (1974) Effects of fire on temperate forests: Northeastern United States.
In: Koslowski TT and Ahlegren EE (eds) Fire and Ecosystems. New York:
Academic Press, pp. 225–250.
Lorimer CG (1993) Causes of oak regeneration problem. In: Loftis DL and
McGee CE (eds) USDA Forest Service General Technical Report SE–84.
Asheville North Carolina: USDA Forest Service, Southeastern Forest
Experimental Station, pp. 14–39.
Luisi N, Lerario P and Vannini A (eds) (1993) Recent advances in studies on
oak decline. Proceedings of an International Congress, 1992. Selva Di
Fassano (Brindisi), Italy. Bari: Universita degli Studi Bari.
Matthews JA and Briffa KR (2005) The ‘Little Ice Age’: Re-evaluation of an
evolving concept. Geografiska Annaler 87: 17–36.
Měřínský Z and Šmerda V (2008a) V mlhách pravěku (Hodonínsko do
příchodu Slovanů). In: Plaček M (ed.) Hodonín: dějiny města do roku
1948. Hodonín, pp. 16–33.
Měřínský Z and Šmerda V (2008b) Svítání středověku (Doba slovanská a
přemyslovských knížat). In: Plaček M (ed.) Hodonín: dějiny města do
roku 1948. Hodonín, pp. 34–54.
Mitchell FJG and Cole E (1998) Reconstruction of long-term successional
dynamics of temperate woodland in Białowieża Forest, Poland. Journal
of Ecology 86: 1042–1059.
Novák V and Pelíšek J (1943) Stručná charakteristika půd na přesypových
pískách v lesní oblasti Dúbrava u Hodonína. Lesnická Práce 8: 225–235.
Pechony O and Shindell DT (2010) Driving forces of global wildfires over
the past millennium and the forthcoming century. Proceedings of the
National Academy of Sciences of the United States of America 107: 19,
167–19,170.
Peel MC, Finlayson BL and McMahon TA (2007) Updated world map of the
Köpen-Gaiger climate classification. Hydrology and Earth System Sci-
ence 11: 1633–1644.
R Development Core Team (2008) A Language and Environmental for Statisti-
cal Computing. Aukland: R Foundation for Statistical Computing.
Rackham O (1988) Trees and woodland in a crowded landscape – The cultural
landscape of the British Isles. In: Birks HH, Birks HJ, Kaland PE et al.
(eds) The Cultural Landscape – Past, Present and Future. Cambridge:
Cambridge University Press, pp. 53–77.
Rackham O (2008) Ancient woodlands: modern threats. New Phytologist 180:
571–586.
Rasband WS and Image JUS (1997–2011) National Institutes of Health,
Bethesda, Maryland, USA, imagej.nih.gov/ij/.
Reille M (1992) Pollen et spores d´Europe et d´Afrique du Nort. Laboratoire
de botanique historique et palynologie. Marseille.
Reille M (1995) Pollen et spores d´Europe et d´Afrique du Nort. Supplement 1.
Laboratoire de botanique historique et palynologie. Marseille.
Reille M (1998) Pollen et spores d´Europe et d´Afrique du Nort. Supplement 2.
Laboratoire de botanique historique et palynologie. Marseille.
Reimer PJ, Baillie MGL, Bard E et al. (2004) IntCal04 terrestrial radiocarbon
age calibration, 0–26 cal kyr BP. Radiocarbon 46: 1029–1058.
Ritchie CJ (1995) Tansley Review No.83. Current trends in studies of long-
term plant community dynamics. New Phytologist 130: 469–494.
Roleček J (2007) Vegetace subkontinentálních doubrav ve střední a východní
Evropě. Dissertation, Masaryk University Brno.
Rybníčková E and Rybníček K (1972) Erste Ergebnisse paläogeobotanische
Untersuchungen des Moores bei Vracov, Südmähren. Folia Geobotanica
Phytotaxonomica 7: 285–308.
Schlachter K and Horn S (2010) Sample preparation methods and replica-
bility in macroscopic charcoal analysis. Journal of Paleolimnology 44:
701–708.
Segerström U (1997) Long-term dynamics of vegetation and disturbance of
southern boreal spruce swamp forest. Journal of Vegetation Science 8:
259–306.
Spitzer L, Konvicka M, Benes J et al. (2008) Does closure of traditionally man-
aged open woodlands threaten epigeic invertebrates? Effects of coppicing
and high deer densities. Biological Conservation 141: 827–837.
Sugita S (1994) Pollen representation of vegetation in Quaternary sedi-
ments. Theory and methods in patchy vegetation. Journal of Ecology
83: 879–898.
Svobodová H (1989) Rekonstrukce přírodního prostředí a osídlení v okolí
Mistřína. Palynologická studie. A reconstruction of natural environment
and settlement in the environs of Mistřín. A palynological study. Památky
archeologické 80: 188–206.
Svobodová H (1990) Vegetace jižní Moravy v druhé polovine prvého tisíciletí.
Archeologické rozhledy 42: 170–205.
Svobodová H (1997) Die Entwicklung der Vegetation in Südmähren
(Tschechien) während des Spätglazials und Holozäns – eine palynolo-
gische Studie. Verhandlungen der Zoologisch -Botanischen Gesellschaft
Österriech 134: 317–356.
Szabó P (2010a) Ancient woodland boundaries in Europe. Journal of Histori-
cal Geography 36: 205–214.
Szabó P (2010b) Driving forces of stability and change in woodland structure:
A case-study from the Czech lowlands. Forest Ecology and Management
259: 650–656.
Tinner W, Condera M, Ammann B et al. (2005) Fire ecology north and south of
the Alps since the last ice age. The Holocene 15: 1214–1226.
Tolasz R, Milíková T, Valeriánová A et al. (2007) Atlas podnebí Česka. Climate
atlas of Czechia. Olomouc: ČHMÚ and Univerzita Palackého.
Van Geel B, Bohncke SJ and Dee H (1980/1981) A palaeoecological study
of an upper Late Glacial and Holocene sequence from ‘de Borchert’, the
Netherlands. Review of Paleobotany Palynology 31: 367–448.

56 The Holocene 23(1)
Van Lear DH (1991) Fire and oak regeneration in the southern Appalachians. In:
Nodvin SC and Waldrop TA (eds) Fire and the Environment: Ecological and
Cultural Perspectives. 20–24 March 1990; Knoxville, Tennessee General
Technical Report SE-69. Asheville, North Carolina: US Department of Agri-
culture, Forest Service, Southeastern Forest Experiment Station, pp. 15–21.
Vera FWM (2000) Grazing Ecology and Forest History. Wallingford & New
York: CABI Publishing.
Veski S, Koppel K and Poska A (2005) Integrated palaeoecological and histori-
cal data in the service of fine-resolution land use and ecological change
assessment during the last 1000 years in Rõuge, southern Estonia. Journal
of Biogeography 32: 1473–1488.
Vodka S, Konvička M and Čížek L (2009) Habitat preferences of oak-
feeding xylophagous beetles in a temperate woodland: Implications
for forest history and management. Journal of Insect Conservation 13:
553–562.
Vykoupil L (2008) Pod svrchovaností převislého rtu (Ve vlastnictví Habsburků
do zrušení patrimoniální správy) 1762–1848. In: Plaček M (ed.) Hodonín:
dějiny města do roku 1948. Hodonín, pp. 260–288.
Watt SA (1919) On the causes of failure of natural regeneration in British
oakwoods. Journal of Ecology 7: 173–203.
Zólyomi B (1957) Der Tatarenahorn-Eichen-Lösswald der zonalen Wald-
steppe. Acta Botanica Academiae Scientiarum Hungaricae 3: 401–424.