Review of Palaeobotany and Palynology 209 (2014) 41–51

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Review of Palaeobotany and Palynology

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Research paper

A 30,000-year pollen record from Mire Kupena, Western Rhodopes

Mountains (south Bulgaria)
Spassimir Tonkov a,⁎, Maria Lazarova b, Elissaveta Bozilova a, Dimiter Ivanov b, Ian Snowball c
a
Laboratory of Palynology, Department of Botany, Faculty of Biology, Sofia University “St. Kliment Ohridski”, Sofia 1164, Bulgaria
b
Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, Sofia 1113, Bulgaria
c
Department of Earth Sciences, Natural Resources and Sustainable Development/Geophysics, Uppsala University, Uppsala SE-752 36, Sweden

a r t i c l e i n f o a b s t r a c t

Article history: A palynological study was performed on a 2 m thick sediment sequence recovered from Mire Kupena (1356 m), a
Received 24 December 2013 former lake in the Western Rhodopes Mountains (south Bulgaria) and supported by radiocarbon dating. The re-
Received in revised form 24 June 2014 cord extends back to ca. 30,000 cal. yrs. BP (Middle Pleniglacial) when the study area was occupied by wooded
Accepted 29 June 2014
steppe composed of Pinus sp., Pinus peuce, some Betula, Juniperus, and cold-tolerant herb vegetation dominated
Available online 9 July 2014
by Artemisia and Chenopodiaceae. In addition, the almost continuous presence of deciduous Quercus, Tilia,
Keywords:
Corylus, Abies, Picea, Fagus, Alnus, and Carpinus betulus pollen suggests interstadial conditions and this area
Pollen analysis could be recognized as a montane refugial place. A reconstruction of the interval between ca. 24,000 and
Vegetation history 15,000 cal. yrs. BP was not possible due to an extremely low sediment accumulation rates or, more likely, a hiatus.
Middle Pleniglacial The lateglacial landscape was dominated by mountain-herb steppe vegetation with isolated stands of Pinus,
Interstadial Betula and shrubland of Juniperus. The afforestation in the Early Holocene started with broad-leaved forests com-
Mire Kupena posed of Quercus with C. betulus, Carpinus orientalis/Ostrya, Ulmus, Tilia and Corylus and minor amounts of Pinus,
Rhodopes Mountains Betula and Abies. In the Late Holocene (bca. 4700 cal. yrs. BP) Fagus began to gain importance chiefly at the ex-
pense of the mixed oak forests, while after ca. 2000 cal. yrs. BP forests of Pinus sylvestris with some P. peuce quickly
expanded around the mire. A comparison with other long palynological records from the mountains and low-
lands of Bulgaria and in northern Greece reveals not only common trends in the vegetation development that
are a reflection of the climate changes, but also site-specific features related to the location and topography of
each site.
© 2014 Elsevier B.V. All rights reserved.

1. Introduction
The location of Bulgaria in the middle part of the Balkan peninsula
creates a high diversity of modern vegetation, topography and climate
conditions, and provides encouraging possibilities for investigations of
past changes in vegetation and climate and the impact of humans. In re-
cent years abundant paleoecological information since the termination
of the last glaciation has been collected from terrestrial sites with
relatively consistent chronological control in the Rila (Bozilova and
Tonkov, 2000, 2011; Tonkov et al., 2008, 2011, 2013a) and northern
Pirin mountains (Tonkov et al., 2002; Stefanova and Ammann, 2003;
Stefanova et al., 2006a,b; Marinova and Tonkov, 2012), and the Thracian
lowlands (Magyari et al., 2008; Connor et al., 2013). These studies re-
vealed the basic stages and trends in vegetation development, climate
fluctuations, possible location of refugia, tree migrations, and patterns
⁎ Corresponding author at: Laboratory of Palynology, Department of Botany, Faculty of
Biology, Sofia University “St. Kliment Ohridski”, 8 Dragan Tzankov blvd., Sofia 1164,
Bulgaria. Tel.: +359 02 8167314; fax: +359 02 8656641.
E-mail address: stonkov@abv.bg (S. Tonkov).
of human impact and they have also enriched our knowledge about
the important role of the Balkans as one of the key areas for the postgla-
cial plant recolonization of Europe (Lang, 1994; Willis, 1994; Bozilova
et al., 1996; Tzedakis, 2004; Jalut et al., 2005; Magri et al., 2006;
Tzedakis, 2009; Fletcher et al., 2010; Tzedakis et al., 2013).
In the context of the above-mentioned palynological and paleoeco-
logical studies in Bulgaria a particular role is assigned to the Rhodopes
Mountains, a vast massif located in the southern part of the country,
with an extension to northern Greece. It is thought that this area was
not glaciated during the Quaternary (Vlaskov, 2002) where a number
of plant species have survived the northern hemisphere glacials. Until
now, indications in support of this hypothesis were obtained from con-
tinuous pollen profiles of lateglacial age collected from mires and peat-
bogs in the Biosphere Reserve Kupena (Bozilova et al., 1989; Huttunen
et al., 1992) and the Shiroka Polyana locality (Filipovitch and Lazarova,
2003; Stefanova et al., 2006a). The influence of the periglacial climate
with low temperatures and insufficient precipitation resulted in the dis-
tribution of herb mountain-steppe vegetation dominated by Artemisia–
Chenopodiaceae–Poaceae and other cold/dry-tolerant species at alti-
tudes between 1000 and 1300 m, and stands of Pinus, Betula with

http://dx.doi.org/10.1016/j.revpalbo.2014.06.002
0034-6667/© 2014 Elsevier B.V. All rights reserved.
42 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51
shrubland of Juniperus–Ephedra. The results suggest that during the
Lateglacial these areas sheltered a number of deciduous (Quercus,
Carpinus betulus, Acer, Fagus, Corylus, etc.) and coniferous (Pinus, Abies,
Picea) trees in places where the local microclimatic conditions favored
their growth and reproduction (Bozilova et al., 2011).
In an attempt to extend the paleoecological evidence from the
Rhodopes Mountains beyond the Lateglacial a new sediment core
from Mire Kupena was collected and the preliminary palynological in-
formation indicated Middle Pleniglacial age (Tonkov et al., 2013b).
This paper presents in more detail the development of the vegetation
and climate changes in the study area during the last ca. 30,000 years
and compares the results with other long pollen profiles from sites in
Bulgaria and northern Greece that are supported by reliable chronolog-
ical frameworks.
2. The study area
2.1. Physico-geographical characteristics and modern vegetation
The Rhodopes Mountains are a large massif, approximately 240 km
W–E and 100 km N–S, which occupy 14,730 km2 in South Bulgaria
and 3270 km2 in Northern Greece. It is divided into the Western
Rhodopes (8732.1 km2 ), with the highest peak Goljam Perelik
(2191 m), which is characterized by vast level stretches that are
most prominent in the interior parts at 1500 m a.s.l., and the Eastern
Rhodopes (1463 m), which is characterized by wide valleys and short ra-
vines. The slightly inclined, typical wide valleys are crossed by meander-
ing rivers and streams. Geologically, the core of the mountains is a large
granite pluton covered by metamorphic rocks (gneisses, schists and mar-
ble) of Paleozoic and pre-Paleozoic ages. The contemporary relief of the
Western Rhodopes was shaped during the Neogene–Quaternary when
the rising vault was broken into large blocks by numerous faults. Some
of these blocks subsided and formed intermontane structural basins
(Galabov et al., 1977). During the Quaternary the highest parts above
2000 m were under the influence of periglacial processes (Vlaskov, 2002).
The climate of the Western Rhodopes is a montane variant of the
Transitional Continental type. The mean annual air temperature is
5 °С–10 °С at 1000 m a.s.l. with mean January temperatures below
0 °С, while the mean July temperature depends on the altitude. The
mean annual precipitation is 600–800 mm with a maximum in May–
June and a minimum in August–September. A Mediterranean influence
is more clearly expressed in the southern parts. The northern slopes of
the mountains are in a precipitation shadow and the amount of annual
precipitation is less than in the interior parts (Velev, 2002). The basic
soil types are brown forest (60%) up to 1700–1900 m, cinnamomic-
forest (up to 800 m), humic-carbonate and mountain-meadow on the
highest parts (Ninov, 1997).
According to the most recent geobotanical division of the country
the Western Rhodopes Mountains are recognized as a separate region
characterized by vast distribution of coniferous forests dominated by
Picea abies (L.) Karst. and Pinus sylvestris L. (Bondev, 2002). The decidu-
ous forests occupy areas at lower altitudes predominantly in the eastern
and northern parts. They are composed mainly of Quercus dalechampii
Ten. with occasional presence of Quercus frainetto Ten. and Quercus
cerris L. on south-facing slopes. Mixed forests of Q. dalechampii, Fagus
sylvatica L., Carpinus betulus L. with participation of Ostrya carpinifolia
Scop. occur in the valleys. Quite often mixed coniferous–deciduous
communities are also found. The present-day vegetation composition
has, in many places, been influenced by grazing herbivores, cultivation
of potatoes and crops, timber production and more recently an increase
in the intensity of touristic activities.
2.2. Study site
Mire Kupena is a former lake (41° 59′ 07.5″ N, 24° 19′ 05.1″ E;
1356 m a.s.l.) found on the territory of the Biosphere Reserve Kupena
in the Western Rhodopes Mountains (Fig. 1). The reserve is located be-
tween 600 and 1400 m a.s.l. on a north-facing slope and occupies an
area of 1086.4 ha. Its present vegetation is rather diverse. The lower
parts are dominated by plant communities of Quercus dalechampii
mixed in some places with Carpinus betulus, Fagus sylvatica, Pinus nigra
Arn. and Pinus sylvestris. Higher up, plant communities of F. sylvatica
dominate, followed by those of P. sylvestris with some Abies alba Mill
and Picea abies. Stands of P. abies are also found. The mire formed in a
depression and covers an area of 6 ha, fed by precipitation and surface
runoff. During spring the water depth can increase to 50 cm due to
rapid snow melt. The mire is surrounded by an almost pure forest of
P. sylvestris with an admixture of P. abies, A. alba, F. sylvatica, Betula
pendula Roth., Sorbus aucuparia L., Q. dalechampii and an undergrowth
of Juniperus communis L., Vaccinium myrtillus L. and Vaccinium vitis
idaea L. The mire vegetation is represented by sparse moss cover of Sphag-
num angustifolium (C. Jens. ex Russ.) C. Jens., Sphagnum platyphyllum
(Lindb. ex Braithw.) Sull. ex Warnst., Sphagnum subsecundum Nees in
Sturm and herbs like Carex riparia Curt., Carex nigra (L). Rchb., Carex
acuta L., Carex echinata Murr., Eriophorum angustifolium Honck., Galium
palustre L., Juncus effusus L., Juncus conglomeratus L., Potentilla erecta (L.)
Rausch., Potentilla palustris (L.) Scop., Ranunculus repens L., etc. During
the last two decades most of the mire hummocks have been overgrown
by pine trees (Huttunen et al., 1992; Tonkov et al., 2013b) (Fig. 2).
3. Material and methods
3.1. Sediment core and lithology
The new core Kupena-3 was collected with a Dachnowsky hand
corer from the peripheral open northeastern part of the mire in 2009
(Fig. 2). The coring was aborted at a depth of 200 cm because it was
not possible to penetrate the underlying strata. The lithology of the
sediments of all three cores from this site is presented in Table 1. The
cores Kupena-1 and Kupena-2 were retrieved from the central part of
the mire about 50 m from each other (Huttunen et al., 1992).
3.2. Pollen analysis
Sampling for pollen analysis was carried out at 5 cm intervals. The
laboratory treatment of the samples followed the standard acetolysis
procedure (Faegri and Iversen, 1989) after removal of siliceous material
with HF acid (Birks and Birks, 1980). The pollen sum (PS) used for
percentage calculations was based on AP (arboreal pollen) + NAP
(non-arboreal pollen), excluding spores of mosses and pteridophytes,
and pollen of aquatics and Cyperaceae. Their presence was expressed
as percentages of the PS. The average number of pollen grains (PS)
counted per sample from the clay/sandy clay sediments (70–200 cm)
was 400–520 and 900–1100 from the uppermost 70 cm of peat and
clay. The identifications of spores and pollen were made using reference
collections, the keys in Beug (2004), Faegri and Iversen (1989) and
Moore et al. (1991). For calculations and construction of the pollen dia-
gram (Fig. 3a and b) the program TGView version 2.0.2 was applied
(Grimm, 2004). The delimitation of the boundaries of the local pollen
assemblage zones (LPAZ) was promoted by CONISS (Grimm, 1987) in
an attempt to correlate the zones with those identified in a previously
studied core from this site, Kupena-2 (Huttunen et al., 1992) (Fig. 4).
3.3. Radiocarbon dating
The radiocarbon age of 6 bulk sediment samples was determined in
the Radiocarbon Dating Laboratory at the University of Lund, Sweden.
The calibration (95.4% probability) was performed with the OxCal v4.1
program (Bronk Ramsey, 2011) and the IntCal13 calibration data
(Reimer et al., 2013). The results and the mean values are shown in
Table 2. The radiocarbon dates from the cores Kupena-1 and Kupena-2
 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51 43

Fig. 1. Map of the Balkans with palynological sites mentioned in text: Western Rhodopes Mountains, Mire Kupena — open triangle, 1 Shiroka Polyana (Filipovitch and Lazarova, 2003;
Stefanova et al., 2006a); Rila Mountains, 2 Lake Ribno and Lake Trilistnika (Tonkov et al., 2008, 2011, 2013a); Pirin Mountains, 3 Lake Kremensko-5 and 4 Lake Bezbog (Stefanova
et al., 2006a,b); 5 Mire Straldzha (Connor et al., 2013); 6 Tenaghi Philippon (Wijmstra, 1969; Müller et al., 2011); and 7 Lake Ioannina (Tzedakis et al., 2002, 2004).

(Huttunen et al., 1992) are also included and their calibrated ages are 4.2. Kupena-3: pollen stratigraphy
presented for the first time.
A brief summary of the LPAZ delimited is outlined as follows (Fig. 3):

4. Results and discussion LPAZ Kup3-1, 200–181 cm (Artemisia–Chenopodiaceae–Pinus

4.1. Radiocarbon chronology and some chronostratigraphical considerations
The two radiocarbon dates LuS 8989 and LuS 9332 from the lower
part of core Kupena-3 confirmed the Middle Pleniglacial age of the
sediments studied. The remainder of the radiocarbon dates refers to
the Lateglacial and the Holocene. The analysis of the calculated accumu-
lation rates reveals a non-linear sedimentation rate in this former lake
basin with a complex topography (Table 2). The first important change
in the lithological composition is observed for the Lateglacial when the
deposition of hard, gray sandy clay ceased and was replaced by clay/
peaty gyttja. At the beginning of the Holocene the sedimentation rate
had been ca. 0.014 cm/yr, whereas after 8300 cal. yrs. BP it has changed
to ca. 0.004 cm/yr. The accumulation of sedge peat started ca. 9300 cal.
yrs. BP. An important consideration established from the pollen diagram
Kupena-3 is that most of the Late Pleniglacial and the transition to the
Lateglacial are not represented, either due to very low sediment
accumulation rates or, more likely, to a hiatus (Tonkov et al., 2013b).
diploxylon-type)
Non-arboreal pollen dominates with 60–80% attributed mainly
to Artemisia (up to 40%), Chenopodiacae (up to 15%) and Poaceae
(5–10%). Values below 5% are recorded for Achillea-type, Dianthus-type,
Scleranthus, Galium-type, Apiaceae, Brassicaceae and Helianthemum. Ar-
boreal pollen is represented by Pinus diploxylon-type (20–35%), Pinus
peuce (up to 4%), Betula, Quercus cerris-type and Juniperus with 2–3%
each. A few pollen grains of Picea, Abies, Corylus, Tilia, Carpinus, Alnus
and Juglans are also found. Characteristic features are the abundance of
microspores of Isoetes (10%) and some Cyperaceae and Polypodiaceae.
LPAZ Kup3-2, 181–143 cm (Pinus–Artemisia–Betula–Quercus)
Arboreal pollen values reach 60% in this zone. High values are re-
corded for Pinus diploxylon-type (40%), accompanied by Pinus peuce
(5–8%), Betula and Quercus robur-type (2–4% each). Pollen grains of
Picea, Abies, Juniperus, Corylus, and Fagus are frequent in all samples. Pol-
len of Ephedra distachya-type is established for the first time. Substantial
44 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51

Fig. 2. A view of Mire Kupena in 2009 with the coring place (photo by V. Bozukov).

changes in the representation of herb pollen are not visible, with the ex- Dianthus-type, Ranunculus-type, Caltha-type, Brassicaceae, Apiaceae,
ception of some decline of Artemisia to 20–30%. Microspores of Isoetes etc. Pollen curves for Cyperaceae, Polypodiaceae, Potamogeton, Typha/
reach maximum values of 20%. Pollen of Typha/Sparganium-type is Sparganium-type and trilete fern spores are recorded. The presence of
also determined. Isoetes microspores becomes sporadical.

LPAZ Kup3-3, 143–113 cm (Artemisia–Chenopodiaceae–Pinus) LPAZ Kup3-4, 113–78 cm (Pinus–Betula–Quercetum mixtum–Fagus)

Arboreal pollen reaches 65–80%, which is composed of Pinus

At the bottom of the zone the arboreal pollen curve reaches its low-
est values of 5–10% and increases to 30–55% at the top of the zone. Pol-
len of Pinus diploxylon-type is ca. 8–10%, Pinus peuce (b5%), Betula,
Juniperus and Quercus robur-type each ca. 5–7%. Pollen of Picea is absent.
Continuous pollen curves of Carpinus betulus and Carpinus orientalis/
Ostrya exist in this zone although with rather low percentages. The
dominant pollen types are Artemisia (40–50%), Chenopodiaceae with a
peak of 30%, while Poaceae keeps constant values below 10%. The vari-
ety of the rest of non-arboreal pollen is represented by Achillea-type,
diploxylon-type (20–25%), Betula, Quercus robur-type and Quercus
cerris-type (ca. 5% each). Peaks are recorded for Abies and Fagus with
7% each. Pollen of other deciduous trees like Ulmus, Corylus, Tilia, and
Carpinus is also present. This zone contains Fraxinus excelsior-type and
Vitis pollen. Herb pollen is represented by Poaceae (up to 10%), Artemisia
(15–10%), some Chenopodiaceae, Achillea-type, Ranunculus-type,
Rumex, Plantago coronopus-type, Brassicaceae, Apiaceae, etc. Micro-
spores of Isoetes disappear from the fossil record. The presence of pollen
of Cyperaceae (up to 20%) and Potamogeton increases.

Table 1 LPAZ Kup3-5, 77–33 cm (Quercetum mixtum–Pinus)

Lithology of the sediment cores from Mire Kupena (Huttunen et al., 1992; Tonkov et al.,
2013b). This zone contains the highest proportions of deciduous tree pollen,
with the main contributions from Quercus robur-type (10%), Tilia (3%),
Core/depth (cm) Lithology
Ulmus (5%) and Betula (6%). There is a reduction in the presence of
Kupena-1 Abies and Fagus. Pollen of Pinus diploxylon-type and Pinus peuce reached
0–40 Peat 30% and 10%, respectively. Among the herb pollen types an increase
40–110 Gyttja
110–125 Clay gyttja
for Poaceae (up to 15%) is established, together with Achillea-type,
125–130 Sand Ranunculus-type, Brassicaceae, Apiaceae, etc. Cyperaceae (up to 30–
130–170 Clay gyttja 40%), Typha/Sparganium-type (up to 10%), Potamogeton and the trilete
Kupena-2
fern spores reach maximum values. The last finds of microspores of
0–85 Peat Isoetes are recorded.
85–135 Peat
135–150 Silty gyttja LPAZ Kup3-6, 33–0 cm (Pinus–Fagus–Abies–Poaceae)
Kupena-3 Arboreal pollen reaches its maximal values, attributed mainly to
0–56 Peat
Pinus diploxylon-type (up to 55%), Pinus peuce (12%), Fagus (15–20%)
56–60 Dark clay
60–70 Peat and Abies (12%). Picea pollen increases to 3% and Carpinus orientalis/
70–90 Dark gray detritus gyttja clay Ostrya-type reaches 5%. Other deciduous trees like Quercus, Ulmus and
90–110 Gray sandy clay Tilia decline. Compared to the previous zone the participation of non-
110–130 Gray detritus gyttja clay arboreal pollen is similar: Poaceae (10%), Plantago coronopus-type,
130–200 Sandy gray clay
Rosaceae, Brassicaceae, etc. Pollen of Chenopodiaceae disappears
 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51 45

Fig. 3. Pollen diagram of Kupena-3.

while Artemisia is sporadically found. A continuous presence of
Triticum-type pollen is recorded. The components of the local vegetation
are represented by Cyperaceae, the trilete fern spores and low propor-
tions for Polypodiaceae, Typha/Sparganium-type and Potamogeton.
4.3. Vegetation development
In accordance with the radiocarbon chronology the lower parts
of the core (LPAZ Kup3-1 and Kup3-2) span a time interval before
30,000 cal. yrs. BP until ca. 24,000 cal. yrs. BP, which means that the pal-
ynological record from this montane area of Bulgaria is the first to be
produced that starts in the Middle Pleniglacial. The reconstruction of
the vegetation cover for this period reveals that the mountain slopes
were covered by sparse forests of Pinus (most probably Pinus
sylvestris/nigra and Pinus peuce) with stands of Betula, and shrubland
of Juniperus–Ephedra on dry sandy places among herb communities
dominated by Artemisia, Chenopodiaceae, Poaceae, Apiaceae, Achillea,
Ranunculus, Dianthus, Galium, and Helianthemum species. At lower alti-
tudes, under more favorable microclimatic and edaphic conditions,
groups of deciduous (Quercus, Carpinus, Tilia, Ulmus, Corylus, Alnus,
Fagus) and coniferous (Abies, Picea) trees thrived. The regional presence
of such a large variety of trees and shrubs confirms that this part of the
Western Rhodopes Mountains served as one of the montane Middle
Pleniglacial refugial places in the Balkans.
Gradually, the forests of Pinus with groups of Betula and Juniperus
started to expand at the expense of the mountain-herb vegetation and
over the course of several millennia (until ca. 24,000 cal. yrs. BP) the ar-
boreal vegetation constituted an important part of the plant cover. The
wider distribution of trees in the study area resulted from climate im-
provement, which can be associated with interstadial conditions. The
extended and improved GICC05 Greenland ice core timescale marks
the start of several interstadials GI-4, GI-3 and GI-2 at 28,900, 27,780
46 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51

Fig. 4. Correlation of Local Pollen Assemblage Zones (LPAZ) established for Mire Kupena, Shiroka Polyana and Lake Ribno (Abbreviations: Ar = Artemisia, Ch = Chenopodiaceae, Po =
Poaceae, Pi = Pinus, Be = Betula, Ju = Juniperus, Pic = Picea, Ab = Abies, Fa = Fagus, Co = Corylus, Qmix = Quercetum mixtum).

and 23,340 cal. yrs. BP, respectively (Blockley et al., 2012). Taking into
account dating uncertainties and likely lags in vegetation response
there is a potential match between the duration of pollen zone Kup3-2
(ca. 28,530–24,650 cal. yrs. BP) and the above interstadials of Greenland
warming. For the first time, therefore, an interstadial episode during
the Middle Pleniglacial has been palynologically recognized from the
Rhodopes area.
The basin itself resembled a shallow lake, which was fed by rainfall
and water from snow melt. The local hydrological conditions favored
the growth of Isoetes lacustris (Lazarova et al., 2012), a species that exists
today in glacial lakes in the northern Pirin Mountains (Stefanova and
Ivanova, 2000), together with ferns, sedges, Potamogeton, Typha and
Sparganium species. Traces of microspores of Isoetes were already found
in the lateglacial section of core Kupena-2 at ca. 14,000 cal. yrs. BP during
the Allerød interstadial, and these were accompanied by other aquatics
such as Myriophyllum spicatum/verticillatum, Myriophyllum alterniflorum,
Potamogeton and Lemna (Huttunen et al., 1992).
The lack of palynological information for the time interval 24,000–
15,000 cal. yrs. BP is most probably a reflection of an important transfor-
mation in the hydrological regime of the lake. A lowering of the water
level, indicated by the sharp decline of Isoetes lacustris and the rest of
the hydro- and hygrophytes, probably resulted from the dry climate
that prevailed during the Late Pleniglacial, including the Last Glacial
Maximum. This dry climate would have contributed to extremely low
sediment accumulation rates, or more likely, to a hiatus until the begin-
ning of the Lateglacial ca. 15,000 cal. yrs. BP (level 140 cm) (Tonkov
et al., 2013b). Congruently with the vegetation reconstruction for the
Balkans presented by Jalut et al. (2005) we suggest that the harsh envi-
ronmental conditions in the Rhodopes Mountains during this period re-
stricted tree vegetation at low and middle elevations to localities that
provided sufficient moisture and that vast areas were dominated by
vegetation of steppic character composed of Artemisia, Chenopodiaceae,
Poaceae and other cold-tolerant species.
The low time resolution of the lateglacial section of ca. 30 cm thick-
ness (LPAZ Kup3-3) does not allow us to reconstruct details of changes
in the vegetation cover. Nevertheless, the general picture of the vegeta-
tion reconstruction in the surroundings of the lake revealed that herb
communities with various representatives of Artemisia, Chenopodiaceae
and Poaceae species, and a number of cold-resistant heliophilous herbs
from Achillea, Cichoriaceae, Dianthus, Lychnis, Galium, Helianthemum
and Apiaceae dominated in the landscape. Isolated stands of Pinus,
Betula and shrubs of Juniperus also existed. At lower elevations groups
 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51 47

Table 2 the time interval 8300–7700 cal. yrs. BP also confirms an enlargement of
Radiocarbon dates from Mire Kupena (Huttunen et al., 1992; Tonkov et al., 2013b). the mixed oak forests which reached their maximal distribution.
14
C lab. Depth 14
C yrs. BP Cal. yrs. BP Material dated Important changes in the vegetation cover are observed after
no. (cm) 4700 cal. yrs. BP. The dynamic processes of tree migrations in the
Kupena-1 study area during the Late Holocene completed with the establishment
Tln 1004 50–70 9000 ± 40 10,243–9938 (10,090) Gyttja of the recent vegetation pattern. First of all, Fagus started to gain impor-
INRNE 100 70–90 9700 ± 150 11,600–10,583 (11,090) Gyttja tance and to some extent Abies and Picea abies, chiefly at the expense of
Tln 1000 100–120 10,700 ± 64 12,853–12,649 (12,750) Gyttja
the mixed oak forests. On drier places Carpinus orientalis/Ostrya became
Kupena-2 an important element. Secondly, the coniferous forests dominated by
Ua 1549 11–12 2070 ± 100 2318–1827 (2075) Peat Pinus sylvestris with some Pinus peuce, Pinus nigra and Abies started
Ua 1550 36–37 6830 ± 110 7928–7499 (7715) Peat
quickly to occupy the area around the mire. In these forests the admix-
Ua 1551 56–57 7560 ± 125 8599–8051 (8325) Peat
Ua 1552 91–92 9320 ± 185 11,168–10,178 (10,675) Peaty gyttja tures of deciduous trees were preferably distributed on south-facing
Ua 1553 116–117 9825 ± 155 11,820–10,730 (11,275) Peaty gyttja slopes. The radiocarbon chronology for the core Kupena-2 (Huttunen
Ua 1554 148–149 11,875 ± 310 14,729–13,123 (13,925) Sandy gyttja et al., 1992) marked precisely the start of Pinus expansion at 2070 cal.
Kupena-3 yrs. BP. In many places fragmented mixed coniferous–deciduous com-
LuS 9328 35–37 4100 ± 50 4822–4445 (4635) Peat munities developed attributed not only to climate change but also to
LuS 9329 52–53 7535 ± 55 8421–8201 (8310) Peat human interference in the natural forest cover. The first convincing
LuS 9331 114–116 10,820 ± 65 12,881–12,584 (12,730) Detritus gyttja signs of human presence are documented by the continuous presence
clay
of Triticum-type pollen since ca. 4700 cal. yrs. BP.
LuS 9791 125–128 12,145 ± 75 14,212–13,781 (14,000) Detritus gyttja
clay
LuS 8989 142–145 20,680 ± 150 25,061–24,293 (24,680) Sandy clay
LuS 9332 161–164 23,730 ± 160 29,070–27,995 (28,530) Sandy clay 4.4. Local and regional comparisons

4.4.1. Montane and lowland sites in Bulgaria

of more moisture demanding deciduous and coniferous trees existed,
such as Quercus, Corylus, Ulmus, Fagus, Carpinus, Juglans, and Abies.
The pollen spectra indicate that a climatic warming occurred soon
after ca. 12,700 cal. yrs. BP when Artemisia and Chenopodiaceae started
to decline, and were replaced by Pinus, Betula, Quercus, Corylus, and
partly from Ulmus, Carpinus and Fagus. The hydro- and hygrophytes
also resumed their presence in and around the lake (Cyperaceae,
Polypodiaceae, ferns with trilete spores, Potamogeton and Isoetes
lacustris). The changes in the pollen stratigraphy in the counterpart
lateglacial section from core Kupena-2 also indicated such a climatic
improvement (Huttunen et al., 1992) (Fig. 4). To summarize, the poor
resolution of data for the lateglacial period does not allow us to clearly
delimit an interstadial/stadial cycle at Kupena.
In the Early Holocene tree communities started to dominate the
landscape (LPAZ Kup3-4, 11,500 cal. yrs. BP–10,000 cal. yrs. BP), partic-
ularly broad-leaved forests of Quercus with Carpinus betulus, Carpinus
orientalis/Ostrya, Ulmus, Tilia and Corylus. The temporary increase of
Fagus and Abies, and the re-appearance of minor quantities of pollen
of Picea, indicate that the climate became milder and moister. On
more open areas at higher elevations a slight expansion of pine commu-
nities (Pinus sylvestris/nigra and Pinus peuce) took place, together with
stands of Betula and Juniperus, which was commensurate with increased
soil development and stabilization. In general, the herbaceous vegeta-
tion became more diverse, particularly enriched with species such as
Rosaceae, Brassicaceae, Apiaceae and Ranunculus. In accordance with
the pollen diagram from Kupena-2 (Huttunen et al., 1992) Artemisia
and Chenopodiaceae species withdrew from the site after ca. 10,670 cal.
yrs. BP. The lake vegetation was composed of hygro- and hydrophytes
from Cyperaceae, Typha, Sparganium, and Potamogeton, but without
Isoetes lacustris. Ferns were also quite common in the surroundings of
the basin.
After ca. 9300 cal. yrs. BP sedge dominated peat began to form at the
edges of the lake and indicates overgrowing and lowering of the water
level. This change in the hydrological regime caused the disappearance
of Isoetes lacustris and the spread of marshy communities, which be-
came dominated by Cyperaceae after 8300 cal. yrs. BP. Since this time,
the composition of the forests has remained nearly the same except
for some reduction of Abies, Fagus, Carpinus orientalis/Ostrya and
Corylus. The share of Tilia and Ulmus in the oak communities has slightly
increased (LPAZ Kup3-4, ~10,000 cal. yrs. BP–4700 cal. yrs. BP). The pal-
ynological evidence from the core Kupena-2 (Huttunen et al., 1992) for
The new pollen record from Mire Kupena can be compared with pal-
ynological sites of similar age or at least with continuous profiles of
Lateglacial age from the mountains and lowlands of Bulgaria. One paleo-
ecological record from the Western Rhodopes Mountains, which dates
to the onset of the Lateglacial originates from the coniferous forest
belt at 1400 m a.s.l. in the Shiroka Polyana locality (Filipovitch and
Lazarova, 2003; Stefanova et al., 2006b) (Fig. 1). Today the site
is peatland on the place of an overgrown floodplain lake and it is
surrounded by forest of Pinus sylvestris and Picea abies with single
trees of Pinus peuce. The thickness of the lateglacial sediments is
70 cm and their age is indicated by a single radiocarbon date of
13,080 ± 55 14C yrs. BP (15,480 cal. yrs. BP) at the bottom part of the se-
quence where silt was deposited. The lateglacial fossil assemblages are
dominated by non-arboreal pollen, attributed mainly to Poaceae (up to
30%), Artemisia, Chenopodiaceae and other cold-tolerant herbs (Fig. 4).
Low frequencies of pollen of trees and shrubs from Pinus diploxylon-
type, Pinus peuce, Betula, Juniperus, Quercus, Corylus, etc. are continuous-
ly recorded throughout the Lateglacial. The pollen stratigraphy from
this site does not allow one to distinguish an interstadial/stadial cycle.
This weakness of the lateglacial palynological subdivision at Shiroka
Polyana is probably a reflection of the location of the site in the
interland of the Western Rhodopes Mountains where the climate
changes during the Lateglacial were moderate.
In the Early Holocene, forests of Pinus expanded between 11,000 and
10,000 cal. yrs. BP with a maximum extension at ca. 10,500 cal. yrs. BP.
Later on, over the course of several centuries Betula and Quercus in-
creased and were firmly established after 8800 cal. yrs. BP, similar to
the area of Kupena (LPAZ Kup3-5). The herb vegetation dominated by
Poaceae and less Artemisia and Chenopodiaceae species persisted until
ca. 9000 cal. yrs. BP, which was 1500 years later compared to the situa-
tion at Kupena (LPAZ Kup3-4). Low humidity was probably the reason
for the continuation of the steppe vegetation into the Early Holocene
in the Rhodopes sites (Stefanova et al., 2006a). Between 8800 and
8300 cal. yrs. BP a slight increase of Abies is recorded synchronously at
Shiroka Polyana and Kupena. About 7800 years ago the mixed Quercus
forests started to retreat, replaced at many places first by Corylus and
subsequently by Abies and Fagus. The last tree to migrate to Shiroka
Polyana after ca. 6000 cal. yrs. BP was Picea abies, while at Kupena this
role was basically assigned to Fagus after 4700 cal. BP (Bozilova et al.,
2011). A common feature in the vegetation development during the
last 2000 years in the Western Rhodopes Mountains was the expansion
of Pinus (mostly Pinus sylvestris).
48 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51
The pollen sequence from Mire Kupena can also be compared to
radiocarbon dated lacustrine pollen records from the Rila and Pirin
mountains. The palynological information archived in the cores collect-
ed from the glacial lakes Trilistnika (2126 m) and Ribno (2184 m) in the
northwestern Rila Mountains (Tonkov et al., 2008, 2011, 2013a) (Fig. 1)
extends beyond 16,000 cal. yrs. BP when a tendency of warming compa-
rable with the paleoclimate isotopic signal from Greenland ice-cores
(NorthGRIP Members, 2004) was established. The exact time of glacier
retreat from the cirques that now form lake basins remains still un-
known, and probably took place around 20,000–18,000 cal. yrs. BP
(Late Pleniglacial). The lateglacial sequences from both glacial lakes
(see Fig. 6 in Tonkov et al., 2011) clearly indicate a stadial/interstadial
cycle, unlike the sites from the Western Rhodopes Mountains, with
wide distribution of cold-resistant herb vegetation dominated by Arte-
misia–Chenopodiaceae and Poaceae, which would be expected during
the Oldest Dryas and Younger Dryas stadials. This vegetation pattern
was partially enriched by stands of Pinus, Betula, and Juniperus–Ephedra
shrubland at higher altitudes and groups of deciduous trees (Quercus,
Corylus, Carpinus) below them during the Bölling/Alleröd interstadial
complex (15,100–12,900 cal. yrs. BP) (Fig. 4). Pollen and plant macro-
fossil studies on lacustrine deposits in the northern Pirin Mountains
(Lake Kremensko-5, 2124 m and Lake Bezbog, 2250 m) also provide a
coherent picture of the lateglacial vegetation development with large
quantities of Artemisia–Chenopodiaceae–Poaceae with Ephedra shrub-
land during the stadials and the spread of Pinus (Pinus peuce, Pinus
diploxylon-type) and Juniperus during the Bölling/Alleröd interstadial
complex due to the more mesic climate conditions (Stefanova et al.,
2006a).
In the Rila and northern Pirin mountains, the afforestation in the
Early Holocene (11,600–7800 cal. yrs. BP) was characterized by pioneer
Betula forests alongside groups of Pinus and Juniperus at high-mid
altitudes and mixed deciduous oak forests with abundant Tilia, Ulmus,
Acer, and later on Corylus below this zone. In the conditions of a
humid climate (7800–5800 cal. yrs. BP) a coniferous belt composed
of Pinus sylvestris, Pinus peuce and Abies was formed. The last tree
immigrants were Fagus after 4300 cal. yrs. BP and Pinus abies after
3400 cal. yrs. BP. These trees experienced several expansion maxima
after 2600 cal. yrs. BP (Tonkov et al., 2002, 2013a) (Fig. 4).
Connor et al. (2013) published a long pollen record of Middle
Pleniglacial age from Bulgaria's former largest lowland inland water
body Mire Straldzha which is located in the northeastern part of the
Thracian plain (Fig. 1). This record can also be compared with the paly-
nological results from Mire Kupena. Despite some uncertainty in the
age–depth model for the pre-Holocene section of the sequence, several
vegetation phases since ca. 37,500 cal. yrs. BP can be recognized (Fig. 5).
An Artemisia-dominated dry/cold steppe phase with patches of xeric
oak woodstands persisted in the lowlands until ca. 17,900 cal. yrs. BP,
and this was succeeded by semi-desert vegetation dominated by
Chenopodiaceae until 10,300 cal. BP. In the area of Kupena during inter-
vals of the Middle Pleniglacial and throughout the Lateglacial and at the
beginning of the Holocene the open steppe herb vegetation was domi-
nated by montane species of Artemisia, Chenopodiaceae and less
Poaceae. Around Mire Straldzha the vegetation during the interval
10,300 to 8900 cal. yrs. BP was relatively open, resembling forests-
steppe with diverse xerophilous and mesophilous herbs, and stands
of Quercus. Gradually, after 8900 cal. yrs. BP this vegetation type
transformed into oak forests with Ulmus, Corylus and Tilia. At Kupena
the mixed oak forests reached their maximal distribution between
8300 and 7700 cal. yrs. BP. The Holocene afforestation around Mire
Straldzha was quite different, as could be expected of a lowland
area experiencing a more arid climate. Betula did not play a promi-
nent role, while Corylus, Ulmus and Quercus increased simultaneous-
ly. The mixed oak woods in the Thracian plain were subjected to
widespread deforestation, burning and grazing after 4000 cal. yrs.
BP (Filipovitch and Stojanova, 1990; Tonkov et al., 2008; Connor
et al., 2013).
4.4.2. Sites in northern Greece
The new pollen core from Mire Kupena shares common features
with long sequences retrieved from northeastern and northwestern
Greece (Tenaghi Philippon and Ioannina, Fig. 1) which vegetation also
responded to the climatic oscillations that occurred throughout the
last glacial. The extent of forest development at each site was influenced
by intra-regional climatic/ecological gradients (Tzedakis et al., 2004).
The nearest site to Kupena is Tenaghi Philippon (40 m a.s.l.), located
in the landlocked Drama plain where the climate is distinctly conti-
nental with colder winters and annual precipitation of 600 mm
(Tzedakis, 2009). The thick deposits archive the vegetation and cli-
mate changes in this coastal area of the northern Aegean Sea for
the last ca. 1.35 million years (Tzedakis et al., 2006). Most pertinent
to the present study are the uppermost 11 m (pollen zones X to Z) of
the pioneer work by Wijmstra (1969) on core TP II, which can be con-
sidered to cover the same time period as the Kupena-3 profile (Fig. 5)
on the basis of the consistent chronology derived from a series of radio-
carbon dates, the oldest one of 28,840 ± 300 14C yrs. BP (30,850 cal. yrs.
BP; see Fig. 2 in Wijmstra, 1969). In summary, the time interval 30,850–
24,000 cal. yrs. BP (subzones X1 to X3) is characterized by the domi-
nance of open Artemisia–Chenopodiceae steppe vegetation with some
increases of Pinus, accompanied by Quercus and Juniperus, during the in-
terstadial (subzone X2). The major part of pollen zone Kup 3-2 features
an enlargement of the pine forests (Pinus sylvestris/nigra, Pinus peuce)
with some Betula and Juniperus at higher altitudes, and Quercus
below them. Since 24,000 cal. yrs. BP until the onset of the Lateglacial
(subzones X4 and X5) the landscape at Tenaghi Philippon was dominat-
ed by open dry Artemisia–Chenopodiaceae vegetation with sporadical
occurrences of Pinus, Juniperus and Betula, as would be expected in a
lowland site.
Part of the new centennial-scale-resolution pollen record from
Tenaghi Philippon (core TP-2005) which covers the last 73,000 years
(MIS 1–4) according to 20 AMS 14C dates, is characterized by a series
of interstadials (short-term expansions of populations of Pinus, decidu-
ous Quercus, Betula, Juniperus) interrupted by dominance of dry steppe
of Artemisia–Chenopodiaceae (see Fig. 3 in Müller et al., 2011). The
spread of tree populations during interstadials was most likely facilitat-
ed by short-term increases of precipitation in otherwise arid environ-
ments of the eastern Mediterranean during the last glacial. Several
interstadials with somewhat higher contribution of Pinus are clearly
recognized at 29,100, 28,100 and 21,900 cal. yrs. BP, and can be linked
with the Greenland interstadials (GIS 4-2) (Müller et al., 2011). Indica-
tions of similar alternations between dry steppe vegetation and tree
expansions are hardly detectable at Kupena. A substantial change in
the vegetation at Tenaghi Philippon occurred after 16,570 cal. yrs. BP
when deciduous and a few evergreen Quercus were quickly established,
accompanied at the beginning of the Holocene by Ulmus and Tilia, while
Pinus was restricted to higher elevations.
The second site with a long pollen archive of the climate changes
during the last glacial that can be compared broadly with the Mire
Kupena is Lake Ioannina (470 m a.s.l.) (Bottema, 1974; Tzedakis,
1993; Tzedakis et al., 2002, 2004; Lawson et al., 2004). This basin, locat-
ed in northwestern Greece, is an intermontane plateau on the western
flank of the Pindus mountain and receives high annual precipitation
values of 1200 mm, which is in contrast to Tenaghi Philippon
(Tzedakis, 2009). The initial palynological study by Bottema (1974)
showed the presence of pollen of Quercus, Abies, Pinus, Betula, Fagus,
Ulmus, Carpinus betulus and other trees during the Last Glacial Maxi-
mum. The most recent record from this site (core I-284) with improved
chronological control revealed that during the Late Pleniglacial the
climatic events were not severe enough to eliminate the existing tree
populations (Tzedakis et al., 2002, 2004). The explanation lies in the
continued moisture availability combined with high topographic vari-
ability that had allowed vertical migrations of tree populations and
their survival in suitable microhabitats (Tzedakis, 2009). The mean
values of arboreal pollen at this site for the time interval ca. 25,000–
 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51
Fig. 5. Comparison of the vegetation history for the last ca. 30,000 years between Mire Kupena, Tenaghi Philippon (Wijmstra, 1969) and Mire Straldzha (Connor et al., 2013) (for abbre-
viations see Fig. 4 with addition of Qu = Quercus, Pist = Pistacia, Fr = Fraxinus).
18,600 cal. yrs. BP, which includes the Last Glacial Maximum, reached
35% (min. 24%; max. 54%) and suggest the presence of deciduous
Quercus, Abies, Pinus, Betula, Fagus, Ulmus, Carpinus betulus, Ostrya/
Carpinus orientalis and Alnus within the Ioannina catchment (Tzedakis,
2004). Of particular interest is the pattern of the vegetation dynamics
during the Lateglacial (Lawson et al., 2004) representing a gradual tran-
sition between a glacial landscape dominated by steppe-grassland with
limited tree presence (deciduous Quercus). This graduality contrasts
with the fluctuations in pollen frequencies of the main tree (Pinus,
Betula) and herb (Artemisia, Chenopodiaceae, Poaceae) taxa recorded
in the Bulgarian high mountains (Tonkov et al., 2011). The effect of cli-
mate changes on the vegetation in Ioannina basin during the Lateglacial
was mitigated by the topographic setting due to a west–east precipita-
tion gradient, thus reducing the potential for deciduous tree growth to
the east of the Pindus mountains (Tzedakis et al., 2004).
5. Conclusions
The new pollen record from Mire Kupena provides, for the first time,
information on the vegetation development, climate changes, tree mi-
grations and establishment in the montane area of Bulgaria during the
last ca. 30,000 years. This information is also compared with evidence
49
from long pollen archives obtained from other sites in the country and
in the Balkans. We reach the following conclusions:
1. During the Middle and the beginning of the Late Pleniglacial
(ca. 30,000–24,000 cal. yrs. BP) wooded-steppe vegetation was
widely distributed in the study area. In addition to the comparatively
well-represented Pinus (Pinus diploxylon-type, Pinus peuce) the re-
cord indicates nearly continuous, though low presence of pollen of
Betula, Juniperus, deciduous Quercus, Tilia, Abies, Picea, Fagus and
Alnus. These data suggest that the area of Kupena in the Western
Rhodopes Mountains could have served as one of the Middle
Pleniglacial refugial places in the Balkans.
2. The interval 24,000–15,000 cal. yrs. BP, including the Last Glacial
Maximum, is evidently not present in the palynological record of
Mire Kupena.
3. The Lateglacial landscape was dominated by mountain-herb vegeta-
tion with isolated stands of Pinus, Betula and shrubland of Juniperus.
At lower altitudes and in microrefugial habitats groups of more
moisture demanding trees survived (Quercus, Ulmus, Corylus, Fagus,
Carpinus, Abies, etc.).
4. The Holocene afforestation started with broad-leaved forests of
Quercus with Carpinus betulus, Carpinus orientalis/Ostrya, Ulmus,
Tilia and Corylus, whereas on more open areas at higher elevations
50 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51
Pinus (Pinus sylvestris/nigra and Pinus peuce) together with stands of
Betula and Juniperus expanded. The mountain-herb vegetation finally
retreated after ca. 10,670 cal. yrs. BP.
5. The last 5000 years featured two important changes in the vegeta-
tion cover, recorded after 4700 and 2000 cal. yrs. BP, respectively.
Firstly, Fagus and to some extent Abies and Picea abies began to gain
importance. Secondly, coniferous forests of Pinus sylvestris with
some Pinus peuce started quickly to expand around the mire and in
many places fragmented mixed coniferous–deciduous communities
formed.
6. Comparisons with other palynological records from the mountains
and lowlands of Bulgaria reveal common trends in the vegetation
development in postglacial time and specific features for each site
dependent on the local topography, vegetation and impact of
regional climate changes.
7. On a regional geographical scale the pollen evidence from Mire
Kupena complements the diverse complex picture of the vegetation
history and climate changes in different parts of the Balkan peninsula
since the Middle Pleniglacial as reconstructed from long paleoeco-
logical records.
Acknowledgments
The research was partly financed through Project BG0034-GAE-
00100-E-V1-EEA FM “Conservation of biodiversity in hot-spots of glacial
relict plants in Bulgaria” by the European Economic Area Financial
Mechanism 2004–2009. The field coring at Mire Kupena was assisted
by Dr. V. Bozukov and B. Tzenov. Prof. P. C. Tzedakis kindly provided
the most recent paleoecological literature on Tenaghi Philippon and
Lake Ioannina. The authors are grateful to both reviewers, Dr. W. Fletcher
and an anonymous colleague, and to the editor Prof. J. Blackford, who
provided useful comments and suggestions to improve the manuscript.
Thanks are due to Dr. A. Tosheva who helped with the drawing of
Figs. 4 and 5.
References
Beug, H.-J., 2004. Leitfaden der Pollenbestimmung für Mitteleuropa und angrenzende
Gebiete. Verlag Dr. Friedrich Pfeil, München.
Birks, H.J.B., Birks, H.H., 1980. Quaternary Palaeoecology. Edward Arnold, London.
Blockley, S.P.E., Lane, C., Hardiman, M., Rasmussen, S., Seierstad, I., Steffensen, J., Svensson,
A., Lotter, A., Turney, C., Ramsey, C., INTIMATE members, 2012. Synchronisation of
palaeoenvironmental records over the last 60,000 years, and an extended INTIMATE1
event stratigraphy to 48,000 b2k. Quat. Sci. Rev. 36, 2–10.
Bondev, I., 2002. Geobotanical division. In: Kopralev, I. (Ed.), Geography of Bulgaria.
ForKom, Sofia, pp. 336–351 (in Bulgarian).
Bottema, S., 1974. Late Quaternary Vegetation History of North-western Greece(Ph.D.
thesis) Rijksuniv, Groningen.
Bozilova, E., Tonkov, S., 2000. Pollen from Lake Sedmo Rilsko reveals southeast European
postglacial vegetation in the highest mountain area of the Balkans. New Phytol. 148,
315–325.
Bozilova, E., Tonkov, S., 2011. 14. Lake Sedmo Rilsko (Bulgaria): Lateglacial vegetation his-
tory. Grana 50 (3), 232–234.
Bozilova, E., Panovska, H., Tonkov, S., 1989. Pollenanalytical investigation in the Kupena
National Reserve, West Rhodopes. Geographica Rhodopica, 1, pp. 186–190.
Bozilova, E., Filipova, M., Filipovich, L., Tonkov, S., 1996. Bulgaria. In: Berglund, B., Birks, J.,
Ralska−Jasiewiczowa, M., Wright, H. (Eds.), Palaeoecological Events During the Last
15000 years. John Wiley & Sons, Chichester, pp. 701–728.
Bozilova, E., Lazarova, M., Tonkov, S., 2011. The postglacial vegetation history of the West-
ern Rhodopes. In: Beron, P. (Ed.), Biodiversity of Bulgaria 4. Biodiversity of Western
Rhodopes (Bulgaria and Greece). Pensoft & National Museum of Natural History,
Sofia, pp. 11–19.
Bronk Ramsey, C., 2011. OxCal Program v4.1. University of Oxford, Radiocarbon Acceler-
ator Unit.
Connor, S.E., Ross, S.A., Sobotkova, A., Herries, A.I.R., Mooney, S.D., Longford, C., Iliev, I.,
2013. Environmental conditions in the SE Balkans since the Last Glacial Maximum
and their influence on the spread of agriculture into Europe. Quat. Sci. Rev. 68,
200–215.
Faegri, K., Iversen, J., 1989. Textbook of Pollen Analysis. John Wiley & Sons, Chichester.
Filipovitch, L., Lazarova, M., 2003. Palynological research in the Shiroka Polyana locality
(Western Rhodopes). Phytol. Balc. 9 (2), 265–274.
Filipovitch, L., Stojanova, V., 1990. Palynological study of the peat bog near Sadovo.
Fitologiya 38, 22–40 (in Bulgarian).
Fletcher, W., Sanchez Goni, M.F., Allen, J.R.M., Cheddadi, R., Combourieu-Nebout, N.,
Huntley, B., Lawson, I., Londeix, L., Magri, D., Margari, V., Müller, U., Naughton, F.,
Novenko, E., Roucoux, K., Tzedakis, P.C., 2010. Millennial-scale variability during the
last glacial in vegetation records from Europe. Quat. Sci. Rev. 29, 2839–2864.
Galabov, Z., Ivanov, I., Penchev, P., Mishev, K., Nedelcheva, V., 1977. Physical Geography of
Bulgaria. Narodna Prosveta, Sofia (in Bulgarian).
Grimm, E., 1987. CONISS: a Fortran 77 Program for stratigraphically constraint cluster
analysis by the method of incremental squares. Comp. Sci. 13, 13–35.
Grimm, E., 2004. TGView (version 2.0.2). Illinois State Museum, Springfield, USA.
Huttunen, A., Huttunen, R.-L., Vasari, Y., Panovska, H., Bozilova, E., 1992. Late-glacial and
Holocene history of flora and vegetation in the Western Rhodopes Mountains,
Bulgaria. Acta Bot. Fenn. 144, 63–80.
Jalut, G., Carrion, J., David, F., Gonzalez-Samperiz, P., Sanchez-Goni, M., Tonkov, S., Willis,
K., 2005. The vegetation around the Mediterranean basin during the last glacial max-
imum and the Holocene climatic optimum. In: Petit-Mair, N., Vrielinck, B. (Eds.), The
Mediterranean basin: the last two climatic extremes. Explanatory notes of the maps
(scale 1/7000000). MMSH and ANDRA, pp. 37–57.
Lang, G., 1994. Quartäre Vegetationsgeschichte Europas. Gustav Fischer Verlag, Jena.
Lawson, I., Frogley, M., Bryant, C., Preece, R., Tzedakis, P., 2004. The Lateglacial and
Holocene environmental history of the Ioannina basin, north-west Greece. Quat.
Sci. Rev. 23, 1599–1625.
Lazarova, M., Ivanov, D., Bozilova, E., Tonkov, S., Snowball, I., 2012. Late Pleistocene and
Holocene history of genus Isoetes L. (Lycopodiophyta) in the Western Rhodopes
Mountains, Bulgaria: new palynological and palaeoecological data. C. R. Acad. Bulg.
Sci. 65 (10), 1405–1410.
Magri, D., Vendramin, G., Comps, B., Dupanloup, I., Geburek, T., Gömöry, D., Latalowa, M.,
Litt, T., Paule, L., Roure, J., Tantau, I., van der Knaap, W.O., Petit, R., de Beaulieu, J.-L.,
2006. A new scenario for the Quaternary history of European beech populations:
palaeobotanical evidence and genetic consequences. New Phytol. 171.
Magyari, E.K., Chapman, J.C., Gaydarska, B., Marinova, E., Deli, T., Huntley, J.P., Allen, J.R.M.,
Huntley, B., 2008. The ‘oriental’ component of the Balkan flora: evidence of ex­pan-
sion into south−east Europe via the Thracian Plain during the last glacial stage. J.
Biogeogr. 35, 865–883.
Marinova, E., Tonkov, S., 2012. Holocene vegetation history of the northwestern Pirin
Mountain (Bulgaria). Plant fossil record from peat−bog Mozgovitsa. C. R. Acad.
Bulg. Sci. 65 (8), 1087–1094.
Moore, P., Webb, J., Collinson, M., 1991. Pollen Analysis, 2nd edition. Blackwell Scientific
Publications, Oxford.
Müller, U., Pross, J., Tzedakis, P.C., Gamble, C., Kotthoff, U., Schmiedl, G., Wulf, S.,
Christanis, K., 2011. The role of climate in the spread of modern humans into
Europe. Quat. Sci. Rev. 30, 273–279.
Ninov, N., 1997. Soils. In: Jordanova, M., Donchev, D. (Eds.), Geography of Bulgaria. 1997,
pp. 225–259 (in Bulgarian).
North Greenland Ice Core Project Members, 2004. High-resolution record of North-
ern Hemisphere climate extending into the last interglacial period. Nature 431,
147–151.
Reimer, P.J., Bard, E., Bayliss, A., Beck, J.W., Blackwell, P.G., Bronk Ramsey, C., Buck, C.E.,
Cheng, H., Edwards, R.L., Friedrich, M., Grootes, P.M., Guilderson, T.P., Haflidason, H.,
Hajdas, I., Hatté, C., Heaton, T.J., Hoffmann, D.L., Hogg, A.G., Hughen, K.A., Kaiser, K.F.,
Kromer, B., Manning, S.W., Niu, M., Reimer, R.W., Richards, D.A., Scott, E.M.,
Southon, J.R., Staff, R.A., Turney, C.S.M., van der Plicht, J., 2013. IntCal13 and Marine13
radiocarbon age calibration curves 0–50,000 years cal BP. Radiocarbon 55,
1869–1887.
Stefanova, I., Ammann, B., 2003. Lateglacial and Holocene vegetation belts in the Pirin
Mountains (southwestern Bulgaria). The Holocene 13 (1), 97–107.
Stefanova, I., Ivanova, D., 2000. On the systematics and history of the Bulgarian represen-
tative of genus Isoetes L. Phytol. Balc. 6 (1), 31–42.
Stefanova, I., Lazarova, M., Wright, H., 2006a. Elevational gradients during the Late-
Glacial/Holocene vegetational transition in southern Bulgaria. Veg. Hist. Archaeobot.
15, 333–343.
Stefanova, I., Atanassova, J., Delcheva, M., Wright, H., 2006b. Chronological framework for
the Late Glacial pollen and macrofossil sequence in the Pirin Mountains, Bulgaria:
Lake Besbog and LakeKremensko-5. The Holocene 16 (6), 877–892.
Tonkov, S., Panovska, H., Possnert, G., Bozilova, E., 2002. The Holocene vegetation his-
tory of Northern Pirin Mountain, southwestern Bulgaria: pollen analysis and ra-
diocarbon dating of a core from Lake Ribno Banderishko. The Holocene 12 (2),
201–210.
Tonkov, S., Bozilova, E., Possnert, G., Velčev, A., 2008. A contribution to the postglacial veg-
etation history of the Rila Mountains, Bulgaria: the pollen record of Lake Trilistnika.
Quat. Int. 190, 58–70.
Tonkov, S., Possnert, G., Bozilova, E., 2011. The Lateglacial in the Rila Mountains (Bulgaria)
re-visited: the pollen record of Lake Ribno (2184 m). Rev. Palaeobot. Palynol. 166
(1–2), 1–11.
Tonkov, S., Bozilova, E., Possnert, G., 2013a. Postglacial vegetation history as recorded
from the subalpine Lake Ribno (NW Rila Mts.), Bulgaria. Cent. Eur. J. Biol. 8 (1),
64–77.
Tonkov, S., Lazarova, M., Bozilova, E., Ivanov, D., Snowball, J., 2013b. 19. Mire Kupena,
Western Rhodopes Mountains (South Bulgaria). Grana 52 (3), 238–240.
Tzedakis, P.C., 1993. Long−term tree populations in northwestern Greece through multi-
ple quaternary climatic cycles. Nature 364, 437–440.
Tzedakis, P.C., 2004. The Balkans as prime glacial refugial territory of European temperate
trees. In: Griffiths, H. (Ed.), Balkan Biodiversity: Pattern and Process in the European
Hotspot. Kluwer Academic Publishers, Dordrecht, pp. 49–68.
Tzedakis, P.C., 2009. Cenozoic climate and vegetation change. In: Woodward, W.
(Ed.), The Physical Geography of the Mediterranean. Oxford University Press,
Oxford, pp. 89–137.
 S. Tonkov et al. / Review of Palaeobotany and Palynology 209 (2014) 41–51
Tzedakis, P.C., Lawson, I.T., Frogley, M.R., Hewitt, G.M., Preece, R.C., 2002. Buffered tree
population changes in a Quaternary refugium: evolutionary implications. Science
297, 2044–2047.
Tzedakis, P.C., Frogley, M.R., Lawson, I.T., Preece, R.C., Cacho, I., de Abreu, L., 2004.
Ecological thresholds and patterns of millennial-scale climate variability: the re-
sponse of vegetation in Greece during the last glacial period. Geology 32 (2),
100–112.
Tzedakis, P.C., Hooghiemstra, H., Pälike, H., 2006. The last 1.35 million years at Tenaghi
Philippon: revised chronostratigraphy and long-term vegetation trends. Quat. Sci.
Rev. 25, 3416–3430.
51
Tzedakis, P.C., Emerson, B.C., Hewitt, G.M., 2013. Cryptic or mystic? Glacial tree refugia in
northern Europe. Trends Ecol. Evol. 28 (12), 696–704.
Velev, S., 2002. Climatic division. In: Kopralev, I. (Ed.), Geography of Bulgaria. ForKom,
Sofia, pp. 155–156 (in Bulgarian).
Vlaskov, V., 2002. Glacial and periglacial morphosculpture. In: Kopralev, I. (Ed.), Geogra-
phy of Bulgaria. ForKom, Sofia, pp. 60–63 (in Bulgarian).
Wijmstra, T.A., 1969. Palynology of the first 30 meters of a 120 m deep section in northern
Greece. Acta Bot. Neerl. 18 (4), 511–527.
Willis, K., 1994. The vegetational history of the Balkans. Quat. Sci. Rev. 13,
769–788.