Vegetation and climate of Anatolia and adjacent regions during the Last Glacial period

Quaternary International 302 (2013) 110e122

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Vegetation and climate of Anatolia and adjacent regions during the Last Glacial period
ur Dog
an b, * Çetin S¸enkul a, Ug a b

Department of Geography, Afyon Kocatepe University, Afyonkarahisar, Turkey Department of Geography, Ankara University, 06100 Sıhhıye, Ankara, Turkey

a r t i c l e i n f o

a b s t r a c t

Article history: Available online 14 April 2012

Reinterpretation of vegetation and climatic conditions of Anatolia and neighboring regions during Last Glacial (30e15 cal ka BP) have been done using a non-quantitative biomization approach based on previously published plant functional types. The results suggest that the climate was cold and humid before w25 cal ka BP and also during the period w23e19 cal ka BP (except East Anatolia). Forest vegetation was 80e90% of the land cover in the northwestern Anatolia and Black Sea coast, and 50e60% along the Mediterranean coast at the same period. During Heinrich Event 2, from w25 to 23 cal ka BP, steppe vegetation dominated the Anatolian region due to the sudden change to very cold and arid conditions. Approximately between 18 and 16 cal ka BP in the Late Glacial, during Heinrich Event 1, a brief interval of cold and arid conditions resulted in weaker steppe vegetation over some part of Anatolia. Around at 15 cal ka BP, suitable climatic conditions for forest vegetation prevailed across a great part of Anatolia, except for Eastern Anatolia and Western Iran, which are highlands. Ó 2012 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction Pollen records are the key proxy for Quaternary palaeovegetation reconstructions. With changes in the pollen percentages assigned to plant functional types (PFT), they provide important information about Quaternary climate cycles (Prentice et al., 1992a,b; De Noblet et al., 1996; Kutzbach et al., 1998; Peyron et al., 1998; Prentice and Webb, 1998; Cheddadi et al., 2001; Sitch et al., 2003; Leroy and Arpe, 2007). PFT-based vegetation and climate models, which cover the Last Glacial Maximum (LGM) onwards conducted on a regional or continental scale mostly for Europe, AfricaeArabia, the Russian Federation, and North America, attracted considerable attention in recent years (Prentice et al., 1992a, 1996; Jolly et al., 1998; Elenga et al., 2000; Tarasov et al., 2000; Davis et al., 2003; Gachet et al., 2003; Robinson et al., 2006; Wu et al., 2007). However, very little modelling has been undertaken for the Anatolian region despite its location at the junction of three continents, climatic zones and ancient human routes. Recently, Cordova et al. (2009) have set the stage for this work by compiling a database for 99 surface pollen samples and 21 charcoal/macrofossils from the region and maps of the modern vegetation zones. * Corresponding author.
an). E-mail address: [email protected] (U. Dog 1040-6182/$ e see front matter Ó 2012 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2012.04.006

A limited number of studies have focused on the conditions of the Last Glacial in Anatolia (see review in Cordova et al., 2009). For example, a study based on the use of pollen analysis for the palaeovegatation reconstruction of Anatolia during the Last Glacial was conducted by Van Zeist and Bottema (1991). Having become a reference for many studies about palaeoenvironmental condi
lu and tions (e.g. Atalay, 1992; Roberts and Wright, 1993; Kuzucuog Roberts, 1998), this study assumes that only the northern coast of Anatolia was covered with forests during the LGM, while the majority of the Mediterranean coastal region, Aegean and Marmara regions, were covered with steppe forests. The remaining areas were covered by steppe and desert steppe formations. Also, this earlier reconstruction suggests that during the LGM and Late Glacial, climatic conditions in Anatolia were generally cold and arid (Van Zeist and Bottema, 1991). However, results obtained from more recent pollen analyses (Caner and Algan, 2002; Mudie et al., 2002, 2007; Emery-Barbier and Thiébault, 2005; Arslanov et al., 2007; Djamali et al., 2008; Litt et al., 2009) and other proxy studies (e.g. Sarıkaya et al., 2008; Fleitmann et al., 2009; Kwiecien
an, 2010, 2011) performed for the reconstruction of et al., 2009; Dog local palaeoenvironmental conditions during the Last Glacial showed that van Zeist and Bottema’s (1991) reconstruction is need of reappraisal. In order to understand the underlying reasons for these inconclusive results and to rethink the existing interpretations of Last

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Glacial (30e15 cal ka BP) pollen records from 12 locations in Anatolia and surrounding regions (Fig. 1) were reinterpreted using a non-quantitative biomization approach. Thus, this study aims to use existing pollen data and previously determined PFT assignments to build a general palaeovegetation and climate reconstruction of Anatolia for the Last Glacial which is the most debated period. A further aim is to demonstrate how global changes are manifest locally. 1.1. Present climate of Anatolia Anatolia is located on the transition zone of different atmospheric circulation systems. However, Anatolian topography has a strong effect over its climate and vegetation. Three types of main air mass that carry moisture to Anatolia were described by Akçar and Schlüchter (2005). Cold and humid air from the polar North Atlantic transported by westerlies mostly produces winter precipitation over Anatolia. Tropical warm and dry air from mid-Atlantic and North Africa with additional moisture from the Mediterranean produces summer precipitation in southern Anatolia. Continental polar air masses transport dry and cold air from Siberia, which condenses on the North Anatolian Mountains after taking up moisture over the Black Sea (Akçar and Schlüchter, 2005). Precipitation over Anatolia is strongly affected by the topography. For this reason, annual precipitation values can range between 300 and 3000 mm (Fig. 2). The Taurus and North Anatolian Mountains play an important role in the distribution of the moisture over the Anatolian Plateau. The high altitudes of these mountain ranges create a natural barrier between coastal and interior regions
lu and Roberts, 1998; Sarıkaya et al., 2009). (Kuzucuog Major fluctuations in the climate of the Mediterranean Sea and borderlands are intimately connected to changes in the thermohaline and atmospheric circulation patterns over the North Atlantic (Melki et al., 2009). On the other hand, the link between North Atlantic and Mediterranean climate is well demonstrated during the Last Glacial by frequent episodes of rapid changes on the Mediterranean palaeoclimatic records (Melki et al., 2009). Studies on millennial-scale climate variability in the Mediterranean Sea

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during the Last Glacial have showed that this region reacted very sensitively to rapid climatic changes in the North Atlantic region related to the Heinrich events (Bond et al., 1992, 1993; Bard et al., 2000; Robinson et al., 2006; Kwiecien et al., 2009). 1.2. Present vegetation of Anatolia Sudden changes in topography, topographic barrier effects, different temperatures and precipitation conditions, etc., strongly affect horizontal and vertical distribution, floristic and formation characters of vegetation cover in Anatolia. Cool-temperate summer green forest zone (Fagus, Alnus etc.,) lies between 0 and 1000 m a.s.l. and is observed in the coastal belt and also in the Northern Anatolian Mountains within the European-Siberian phytogeograpic region, characterised with humid and temperate climate conditions (Fig. 3). Boreal evergreen conifer forest (Pinus sylvestris, Picea, Abies etc.) occupies coldehumid areas between 1000 and 2000 m. Warm-temperate conifer (Pinus brutia, Pinus pinea) and warm temperate summer/evergreen forest (Quercus coccifera, Quercus cerris etc.,) zone develop under the Mediterranean climate conditions between 0 and 1200 m a.s.l. over mountainous areas behind the Aegean and the Mediterranean coastal belt, in the Mediterranean phytogeographic region. Eurythermic conifer forest (Cedrus, Pinus nigra etc.) can be seen between 1200 and 2200 m in the Irano-Turanian and Mediterranean phytogeographic transition zone. P. nigra and Quercus forest (Quercus robur, Q. cerris etc.) occupy highland areas in the Eastern Anatolia and Central Anatolian Plateau, which are situated in the Irano-Turanian phytogeographic region. 2. Methods The pollen records of 12 locations in Anatolia and surrounding regions, which date back to the Last Glacial period, were analyzed by initially dividing them into three phytogeographic regions: the EuropeaneSiberian, Irano-Turanian, and Mediterranean (Fig. 1, Table 1). Two of the given locations were in the transition zone between the Irano-Turanian and Mediterranean phytogeographic

Fig. 1. Pollen record sites older than 15,000 cal ka BP: 1. Western Black Sea, 2. Marmara Sea 1, 3. Marmara Sea 2, 4. Eastern Black Sea, 5. Eski Acıgöl, 6. Lake Van, 7. Lake Urmia, 8.
ı, 9. Lake Beys¸ehir, 10. Lake Sög
üt, 11. Öküzini 12. Ghab. Present phytogeographic regions: A. European-Siberian phytogeograpic region, B. Irano-Turanian phyKaramık Bataklıg togeograpic region, C. Irano-Turanian and Mediterranean phytogeographic transition zone, D. Mediterranean phytogeograpic region.

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Fig. 2. Annual average precipitation of Turkey (Çiçek and Ataol, 2009).

regions. Arboreal Pollen (AP) species common to all locations are Pinus, Abies, Castanea, Cedrus, Alnus, Fagus, Betula, Acer, Ulmus, Tamarix, Juniperus, Quercus, Olea and Pistacia. However, there are also subspecies of Pinus and Quercus which have different requirements in terms of temperature and precipitation. P. brutia, Q. coccifera, Q. cerris and Quercus aucheri, which prefer temperature and semi-arid conditions, characterize the Mediterranean phytogeographic region. Pinus sylvestris, P. nigra, Quercus macranthera,

Quercus frainetto, Q. robur, Quercus petraea and Quercus hartwissiana, which prefer cool and sub-humid/humid conditions, are found in the European-Siberian and Irano-Turanian phytogeographic regions. Selected Non-Arboreal Pollen (NAP) taxa are Artemisia, Chenopodiaceae, Poaceae, Cyperaceae, Matricaria, and Senecioare. The pollen records were then analyzed by considering seven species (Table 2) that were common and widespread and characterized different temperature and precipitation conditions as

Fig. 3. Modern biomes/ecoregions and cultivated areas of Turkey.

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Table 1 Selected pollen record locations in Anatolia and adjacent regions. No

Location

Elevation (m a.s.l.)

14

1 2 3 4 5 6 7 8 9 10 11 12

Batı Karadeniz Marmara Denizi 1 Marmara Denizi 2 Dziguta Eski Acıgöl Lake Van Lake Urmia
ı Karamık Bataklıg Lake Beys¸ehir
üt Lake Sög Öküzini Ghab

0 0 0 120 1300 1650 1268 1000 1120 1400 305 400

15,380 15,930 21,950 28,930 15,000 20,000 190,000 20,000 16,000 25,000 24,180 47,000

C age BP

Cal.

14

18,456 18,846 26,350 33,452 18,250 23,927 200,000 23,927 19,148 29,997 28,953 50,577

indicated by using a plant functional type-based biomization approach (Prentice et al., 1992a, 2000; Prentice and Webb, 1998; Cramer, 2002) and interpreted similarly to numerical model studies (Olson et al., 1983; Prentice and Sykes, 1995; Haxeltine et al., 1996; De Noblet et al., 1996; Kutzbach et al., 1998; Cheddadi et al., 2001; Otto et al., 2002; Sitch et al., 2003). However, reinterpretation of vegetation and climatic conditions of Anatolia and its surrounding region during the Last Glacial (30e15 cal ka BP) have been made by using a non-quantitative biomization approach based on plant functional types from previously published palynological records (Table 1). The distinctive feature of the approach adopted by this study is that it aims to identify the changes in the climate conditions and the vegetation pattern in Anatolia by inferring the percentages of taxa and their corresponding climatic conditions. Characteristic of cold environments (Tc, min
35  C), Pinus, Artemisia and Chenopodiaceae are the natural cosmopolitan species of Anatolia and are found in the EuropeaneSiberian, IranoTuranian, Mediterranean phytogeographic regions and in the IranoTuranianeMediterranean transition zone. Despite similarities in robustness in terms of summer drought, Pinus and Artemisia are more widespread than Chenopodiaceae in areas where winter precipitation is higher (Singh et al., 1973; El-Moslimany, 1990). The traditional interpretation of Artemisia as an index of semi-arid conditions has also been challenged by Prentice et al. (1992b). Alnus and Fagus (Tc, min
15  C) are woody trees and shrubs characteristic of cool and humid environments (Günal, 1997). Belonging to the EuropeaneSiberian phytogeographic region, both species grow in Northern Anatolia today, particularly on northfacing mountain slopes and humid areas. Quercus (especially Q. coccifera, Q. cerris, Q. aucheri) and Pistacia (Tc, min þ5  C) prefer warmer conditions but can also tolerate lower winter temperatures Table 2 PFT and bioclimatic tolerance values that characterize environmental conditions in Anatolia (Tc; temperature of the coldest month, a; moisture index, degrees Celsius) (Prentice et al., 1996; Peyron et al., 1998; Elenga et al., 2000; Tarasov et al., 2000; Gachet et al., 2003; Leroy and Arpe, 2007). Pollen taxa included

PFT

Climate type

Tc, min ( C)

Tc, max ( C)

min a

Pinus

Boreal evergreen conifer Steppe forb Steppe forb Boreal summer green Cool-temperate summer green Warm-temperate sclerophyll Warm-temperate sclerophyll

Cold


35


2

0.65

Cold Cold Cool


35
35
15

0.65

Cool


15

0.65

Artemisia Chenopodiaceae Alnus Fagus Quercus Pistacia

Warm

5

Warm

5

15.5

C age BP

0.65 0.28

Phytogeograpic regions

References

European-Siberian European-Siberian European-Siberian European-Siberian Irano-Turanian Irano-Turanian Irano-Turanian Transition zone Transition zone Mediterranean Mediterranean Mediterranean

Atanassova, 2005 Mudie et al., 2002 Mudie et al., 2007 Arslanov, et al., 2007 Woldring, 2001 Litt et al., 2009 Djamali et al., 2008 Van Zeist et al., 1975 Bottema and Woldring, 1984 Van Zeist et al., 1975 Emery-Barbier and Thiébault, 2005 Niklewski and van Zeist, 1970

(Prentice et al., 1996). These two species are used as warm environment indicators in vegetation and climate reconstructions (Rossignol-Strick, 1993, 1995, 1999).
üt, Urmia lakes, and Karamık The pollen data from Ghab, Sög
ı (swamp) have been obtained from European Pollen Bataklıg Database (EPD) together with their 14C ages. The pollen diagrams were constructed by using TILIA and TILIAGRAPH computer package (Grimm, 1991) and the 14C chronology of pollen diagrams have been obtained on the basis of pollen 14C ages. Lastly, to ensure the coherence of Pollen diagrams of chosen species, all 14C ages have been converted to cal 14C ages by using the CalPal programme (Danzeglocke, 2010). In addition, these dates have been matched with the chronology of the high-resolution oxygen isotope records from Greenland (Dansgaard et al., 1993) to demonstrate the rate of change among local functional plant types. 3. Reinterpretation of the Last Glacial pollen records in Anatolia and vicinity The studies on pollen analysis so far have assessed only the vegetation pattern of the area and vicinity where the pollen data were obtained. However, they have not investigated the presence of a potential connection between the observed local changes and previously experienced global climate changes. Therefore, the distinctive reinterpretation made in this study derives from the fact that it aims to identify both the vegetation pattern of the area where pollen data were obtained and the reaction given by this local vegetation pattern to the global climate changes. In addition, development of the necessary scientific infrastructure to a large extent to assess the available pollen data recordings more properly has made it necessary to re-examine them. 3.1. EuropeaneSiberian phytogeographic region Pollen data obtained from the western Black Sea (XK-120 near the Turkish-Bulgarian border) (Fig. 1) (Atanassova, 2005) begins at early Late Glacial (18,456 cal BP). The prevailing type of tree in the PFT during the Late Glacial period (between w18.5 and 15 cal ka BP) was Pinus, which had a percentage value w10e20% suggesting a cold environment, whilst pollen types reflecting cool and/or warm environmental conditions (e.g., Fagus, Alnus, Quercus and Pistacia) were rare or nonexistent. Herbaceous or NAP prevailed, comprising mainly Artemisia, Chenopodiaceae and Poaceae which reached 80% (Fig. 4), suggesting cold and arid conditions for this region. As a result, NAP species were more common than AP species. However, decreasing NAP from 16 cal ka BP onwards was replaced by approximately 15% Pinus. In addition, the emergence of Fagus, Alnus and Quercus during this period reveals the presence of

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Fig. 4. PFT and AP/NAP percentage in the Western Black Sea (redrawn from Atanassova, 2005), Marmara 1 and 2 (redrawn from Mudie et al., 2002, 2007) and Eastern Black Sea (redrawn from Arslanov et al., 2007), Eski Acıgöl (redrawn from Woldring, 2001) and Lake Van (redrawn from Litt et al., 2009) locations. Greenland ice core chronology redrawn from Dansgaard et al. (1993) for time correlation.

small pockets in the region. This may have been due to the increase in precipitation despite the still cold climate since 16 cal ka BP. Pollen data from the Marmara Sea 2 (Mudie et al., 2007) (Core Mar-94-5) show that the AP values before w25 cal ka BP were around w80%, with the dominant AP taxa being Pinus (w45%), Quercus (15%), Fagus (5%) and Alnus (5%). Artemisia (10%) and Chenopodiaceae (5%) composed most of the NAP (20%). This aspect of the vegetation pattern shows that relatively cool and humid climatic conditions prevailed throughout the region before w25 cal ka BP. Vegetation cover around the Marmara Sea around w24e23 cal ka BP was reflected in the pollen records. In this period, the NAP percentage reached 40%. Artemisia-Chenopodiaceae percentage within the NAP reached 30% in relation to the previous period. Pollen data from the Marmara Sea 1 (Mudie et al., 2007) (Core Mar-97-11) show that the percentage of Pinus reached 80% within 60e70% AP by between 23 and 18 cal ka BP. Artemisia and Chenopodiaceae percentages did not exceed 20% within the NAP (30e40%) by that time (Fig. 4). This suggests the existence of dense
et al., 1975). Similarly, increased forests in the region (Aytug percentage of pollen values of Alnus (5%) around the same time also suggests cool and humid climatic conditions throughout the region. Furthermore, existence of other taxa, such as Castanea, Cedrus, and Abies (Caner and Algan, 2002; Mudie et al., 2002, 2007), shows that LGM climate conditions in the Marmara region were suitable for a forest ecosystem. Around the Marmara Sea 1, a decrease in AP percentages occurred between w18.2 and 16 cal ka BP and Pinus decreased from 80% to 30e40% (Fig. 4). However, because of the fact that Quercus is more tolerant than Pinus in terms of arid conditions, Quercus started to exist in the region. This reflects a decrease in precipitation rather than a temperature change. On the other hand, within NAP, the percentage of Artemisia and Chenopodiaceae in the vegetation pattern reached 30%. Between 16 and 15 cal ka BP, AP and NAP percentages fluctuated around the Marmara Sea.

According to the pollen data obtained from Dziguta in the eastern Black Sea (near the Turkish-Georgian border) (Arslanov et al., 2007), AP was around 90% before w25 cal ka BP. Pinus, which is a characteristic of cold environments, reached 80% in the AP (Fig. 4). The low percentages of Fagus, Alnus, and Quercus trees within the AP values points to low temperature, and less than 10% Artemisia and Chenopodiaceae in NAP points to high precipitation. This vegetation pattern changed notably between w25 and 23 cal ka BP. During this period (w1.5e2 ka) AP values decreased to w30e40%. The dominant species of NAP, Cyperaceae, Cichortaceae, Asteraceae and Erigeron constituted approximately 40% of the vegetation pattern. Therefore, the climate is interpreted as being cold and arid (Arslanov et al., 2007). After this period, between w23 and 19 cal ka BP, AP values attained 90% Pinus, similar to the preceding period. Therefore, cold and more humid climatic conditions became established in the region. At around w18 cal ka BP, AP values attained 90e95%, comprising of Abies (40%) which characterizes cool and humid environment, became the dominant tree species in the forest ecosystem. The percentage of Fagus and Alnus exceeded w30%, while those of Pinus, Artemisia and Chenopodiaceae exceeded 20% (Fig. 4) (Arslanov et al., 2007). The very low percentage of Quercus may suggest that no significant change occurred in precipitation or temperature. As a result, the coldehumid conditions at Dziguta gave way to cool-humid conditions during the Late Glacial period (18e15 cal ka BP). 3.2. Irano-Turanian phytogeograpic region Pollen data obtained from the Eski Acıgöl (Woldring, 2001), located in Central Anatolia and parts of the Irano-Turanian phytogeographic region, do not include the LGM. Around 18 cal ka BP, the pollen data from Eski Acıgöl shows that NAP (90%) included approximately 40% Artemisia and Chenopodiaceae (Fig. 4). The dominant AP taxa whose total relative abundance was 15% were Quercus, Pistacia and Pinus. NAP decreased to w80% at around 16 cal

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ka BP and the percentage of Artemisia and Chenopodiaceae decreased to w30%. On the other hand, AP values consisting of Quercus and Pinus increased to w20%. This suggests that climate condition for central Anatolia shifted from coldearid to cool-humid towards 18 to 16 cal ka BP. On the basis of the pollen data from Lake Urmia (Djamali et al., 2008, core BH2) w150 km southeast of Lake Van, AP was around 10% before 25 cal ka BP. Chenopodiaceae w20%, Poaceae w30% and Artemisia w40% were dominant in NAP. Vegetation cover in Lake Urmia around w24e23 cal ka BP was reflected in the pollen records. In this period, the NAP percentage was 90%. ArtemisiaChenopodiaceae percentage within the NAP reached the 80% compared to the previous period. Pollen data from Lake Van shows that NAP (95%) species were dominant at 20 cal ka BP (Litt et al., 2009) (Fig. 4). In the vegetation pattern from this date to the Holocene, Chenopodiaceae reached w40% and Artemisia w10%. This data reveals that the Eastern Anatolia Plateau with a mean elevation of 2200 m was cold and arid during the Last Glacial. Pinus exist within AP around Lake Van in the period of 16 cal ka BP. On the other hand, Quercus are seen around Lake Urmia in the same period. These differences within AP can be explained, as the elevation of Lake Urmia is lower than Lake Van as w400 m. AP percentage started to increase during Early Holocene as observed in Van and Urmia (Landmann et al., 1996; Djamali et al., 2008; Litt et al., 2009). 3.3. Irano-Turanian and Mediterranean phytogeographic transition zone
ı (located between Central Pollen data from Karamık Bataklıg and Southwest Anatolia) (Van Zeist et al., 1975) shows that NAP dominated the overall composition of the vegetation around 24 cal ka BP (Fig. 4), reflecting a decrease in temperature and precipitation. The percentage of AP between w23 and 19 cal ka BP increased to 40e50%, with Pinus reaching 15% and Quercus 5e10% (Fig. 5). Within the NAP, Artemisia had a percentage of 40-20%

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and Chenopodiaceae 10e25%. Other species in the environment during this period were Poaceae and Matricaria within NAP, and Cedrus and Betula within AP (Van Zeist et al., 1975). This vegetation composition shows that cold and humid conditions were dominant during the LGM.
ı, a decrease in AP percentages was recorded In Karamık Bataklıg between w18e16 cal ka BP and total NAP pollen types were 60%. NAP percentages decreased to 30% at around 15 cal ka BP and AP values reached 70% (Fig. 5). In the vegetation composition, the dominant pollen taxon Pinus increased to 20%, followed by Quercus. Additionally, Cedrus, Abies and Betula were the other AP pollen types in the environment (Van Zeist et al., 1975). This change in the vegetation shows that the coldehumid conditions dominant in the region during the LGM gave way to a cool-humid environment in the Late Glacial period between 16 and 15 cal ka BP. Pollen data from Lake Beys¸ehir, also located in the transition zone between the Irano-Turanian and Mediterranean phytogeographic regions (Fig. 1) (Bottema and Woldring, 1984) shows that AP values at 19 cal ka BP were between 10 and 15%, comprising Pinus and Cedrus (Fig. 5). NAP (85e90%) comprised of marsh Matricaria-Senecio, Artemisia and Chenopodiaceae totalling 5e15%. This vegetation pattern reveals that Lake Beys¸ehir (perhaps due to its location) was cold and arid in the LGM. Around Lake Beys¸ehir, AP values (10e15%) remained almost the same between 18 and 16 cal ka BP. The dominant AP taxon was Pinus, followed by Quercus. Within the NAP, Poaceae and Senecio remained as the dominants (Bottema and Woldring, 1984). The percentage of Artemisia and Chenopodiaceae remained at 10e15%. An increase in AP values was registered during the interstadials of Late Glacial period. This data shows that the climate was cold and arid around Beys¸ehir during the Last Glacial. 3.4. Mediterranean phytogeographic region
üt, which lies at 1400 m a.s.l. in The pollen data from Lake Sög southwestern Anatolia (Van Zeist et al., 1975), shows that the NAP


ı (redrawn from Van Zeist et al., 1975), Lake Beys¸ehir (redrawn Bottema and Woldring, 1984), Lake Sög
üt (redrawn from Fig. 5. PFT and AP/NAP percentages in the Karamık Bataklıg Van Zeist et al., 1975), Ghab (redrawn from Niklewski and van Zeist, 1970) and Öküzini (redrawn from Emery-Barbier and Thiébault, 2005) locations.

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percentage reached 70e80% between w24 and 23 cal ka BP, and the total percentage of Artemisia and Chenopodiaceae rose to w50% (Fig. 5). The dominant species within the AP was Pinus (w20%). Therefore, the climate was cold and arid during this period. Between 22 and 19 cal ka BP, AP increased to 55% and the dominant species became Quercus (40%). The NAP percentage decreased to 45%, composed largely of Artemisia and Chenopodiaceae. This data suggests that precipitation and temperature in the region became higher. Between 19 and 15.5 cal ka BP, AP percentages around Lake
üt decreased compared to the LGM and further decreased to Sög 30e40% when Pinus and Quercus became the dominant tree species. Other pollen types recorded as Alnus, Cedrus, Betula and Juniperus (Van Zeist et al., 1975). On the other hand, the total percentage of Artemisia and Chenopodiaceae within the NAP was around 20e15%. This data shows that precipitation during the Late Glacial (19e15.5 cal ka BP) was lower in the region compared to the LGM. During the interstadials of the Late Glacial period around at 15 cal ka BP, there was an increase in AP species. Pollen records from the Ghab coring site in the Eastern Mediterranean region (Niklewski and van Zeist, 1970) show that the AP values before w25 cal ka BP were around 60e70%, with the dominant taxa being Quercus (40%), Pinus (10%), Alnus (5%) and Pistacia (5%). Artemisia (10%) and Chenopodiaceae (20%) composed most of the NAP (30%). This aspect of the vegetation pattern shows that relatively warm and humid climatic conditions prevailed throughout the region before w25 cal ka BP. A marked change in the environmental conditions in Ghab around between w24 and 23 cal ka BP was reflected in the pollen records. In this period, the NAP percentage reached its maximum value, and the Artemisia-Chenopodiaceae percentage of the NAP (60%) doubled compared to the previous period (Fig. 5). This shortlived and sudden change in the vegetation pattern reflects a cold and arid environment. After this cold and arid period, AP increased quickly to w60% between 23 and 18 cal ka BP, comprising of Quercus, Pinus and Cedrus (Niklewski and van Zeist, 1970) and the vegetation composition and climatic conditions returned to the similar values as before w25 cal ka BP. Between w18e15.5 cal ka BP, AP values at Ghab showed decreases of Quercus to 20% and Pinus to 5%, while NAP increased to 65%, including Artemisia (20%) and Chenopodiaceae (25%). This vegetation pattern shows that cold and arid climate conditions returned during the early Late Glacial. Pollen data from the Öküzini site, located near Antalya on the Mediterranean coast (Emery-Barbier and Thiébault, 2005), shows that the NAP percentage reached 80% between w25 and 23.5 cal ka BP. The percentage of Quercus within the AP for the period w25 cal ka BP was around 20% (Fig. 5). At the same time, the Mediterranean species of Pistacia, Olea, Acer, Ulmus and Tamarix were also present, though rare (Emery-Barbier and Thiébault, 2005). Within the 80% NAP, the percentage of Artemisia and Chenopodiaceae was w30%. This steppe vegetation pattern characterizes cool and arid environment during this period. The decrease in the percentage of Artemisia after w23.5 cal ka BP and the increase in the percentage values of Pistacia and Olea indicate an improvement in temperature and humidity. Between w16.5 and 15.5 cal ka BP, Artemisia and Chenopodiaceae declined to 10% of the total NAP, which was around 70e75%. However, the percentage of Lactucae and Poaceae increased to 40e50% (EmeryBarbier and Thiébault, 2005). Within the AP, around 20e25%, Quercus decreased to 10%. As a result, similar to other locations in the Mediterranean, the climate around Öküzini Cave was cool and arid around w17e15.5 cal ka BP. However, unlike the other


üt and Ghab), decrease in the AP percentages in this locations (Sög region was remarkable for the same period. 4. Results: the vegetation and climate of Anatolia in the Last Glacial 4.1. Before 25 cal ka BP In this period, a diversity of vegetation types and climates existed throughout Anatolia. In the Marmara region and along the Eastern Black Sea coast, a dense forest cover developed in the cold and humid conditions. During the same period, in the Mediterranean phytogeographic region, there was a forest that covered approximately 50e60% of the land, consisting of cold and cool environment species. Therefore, a widespread forest cover developed in the cold/ cool and humid conditions before w25 cal ka BP around the Marmara Sea, the entire Black Sea coast and Southern Anatolia. 4.2. Between 25 and 23 cal ka BP This period was marked in the Black Sea and Marmara regions by a drastic decrease in AP percentage but a large increase in NAP values. Species characterizing cold and arid environments were dominant in both AP and NAP. Unfortunately, there is no on-site data for this period in the Central and Eastern Anatolian regions, where a continental climate currently exists. However, it can be suggested that steppe vegetation developed under cold and arid climate conditions in this region. While the increase was seen in
üt, NAP species which characterizes cold and arid climate in Sög Ghab and Öküzini belonged to Mediterranean phytogeographic region in this period, a significant decrease was seen in AP percentages. These cold and arid climatic conditions may be a reflection of a Heinrich Event (H2) effect in some parts of Anatolia. 4.3. Between 23 and 19 cal ka BP In this period, climatic conditions and vegetation patterns of Anatolian displayed similar characteristics as before ca. 25 cal ka BP (Fig. 6). During this time, humid and cold conditions became dominant around the Marmara Sea and the Eastern Black Sea region. In addition, under the cold/arid climate conditions of the TurkisheBulgarian border region and the Black Sea shores of Bulgaria, vegetation represented by NAP kept its dominant character, similar to Central Europe (Willis and Andel, 2004). On the other hand, coldehumid climatic conditions can be inferred for the Mediterranean and Central Anatolia transition zone. Eastern Anatolia and Northwestern Iran were dominated by cold and semi-arid conditions. Unfortunately, pollen data for this period is not available for Central Anatolia. 4.4. Between 19 and 15 cal ka BP While cold and partially arid climatic conditions were dominant in the western Black Sea region (around TurkeyeBulgaria border), there was a decrease in AP around the Marmara Sea and Southwestern Anatolia. This incident in the Late Glacial (18e15.5 cal ka BP) may be attributed to the effect of Heinrich Event 1 (H1), although the Eastern Black Sea region was not clearly affected by this event (Fig. 7). The decrease in precipitation and temperature conditions in the eastern Mediterranean region during H1 (between 17 and 15.5 cal ka BP) caused a notable decrease in AP values. Eastern Anatolia was cold and arid during the Late Glacial. Probably, the effects of the Siberian High Pressure systems rather than Mediterranean systems must have delayed the adaptation of plant cover to post-LGM climate change.

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Fig. 6. PFT types in vegetation communities and environmental conditions during the LGM (around 20e19 cal ka BP) in Anatolia and adjacent regions. The borders of pluvial lakes adapted from Erol (1979) and Kashima (2002). Glaciated and permanent snow areas have been adapted from Erinç (1978, 2001), Çiçek et al. (2003) and Çiner (2003), and appointed permanent snow line from Erinç (1978).

AP values increased at some sites (Marmara, Dziguta, Eski Acıgöl, Karamık, Ghab) in Anatolia around at 15 cal ka BP. AP species suggest that cool or warm and humid environments started to become dominant in these sites. As a result, it may be suggested that warmer and wetter climatic conditions were conducive to forest growth largely throughout Anatolia in the Bølling/Allerød interstadials in the Late Glacial.

5. Discussion 5.1. Environmental conditions before 25 cal ka BP In their interpretations of pollen records for this period, Arslanov et al. (2007) claim that the Eastern Black Sea region was arid, although Mudie et al. (2007) reported relatively mild, wet

Fig. 7. PFT types in vegetation communities during the Late Glacial (w18 cal ka BP) in Anatolia and adjacent regions.

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conditions with winter temperatures not much lower than today for the Marmara region. Mudie et al. (2007) showed that the summer sea surface temperature in the Marmara Sea was almost 10  C lower than now. However, some high values of Pinus in AP may suggest that the climate in the region before w25 cal ka BP was cool and humid. Similarly, Niklewski and van Zeist (1970) concluded from the pollen records of Ghab (in the eastern Mediterranean region) that the climate was cold and arid despite forest cover of approximately 70% in this period. However, the PFT assessment and vegetation pattern reports before w25.1 cal ka BP show that the region was cold and humid. The increased lake levels in Anatolia and vicinity are harmonious with the results of pollen data which are limited in number for this period. From prior to w26.5 to 24.6 cal ka BP, the Palaeo-Lake Konya reached a high level (20 m depth), covering the entire Konya plain and inundating the limestone bench in the middle of the plain (Fontugne et al., 1999). Similarly, the Lake Van level reached a highstand between 26 and
lu et al., 2010). 24.5 cal ka BP (Kuzucuog

synchronous with H2. In the Anatolia, the contraction of PalaeoLake Konya between 24.6 and 23.2 cal ka BP (Fontugne et al., 1999)
lu et al., 2010) and Lake Van between 24 and 22 cal ka BP (Kuzucuog also occurred during H2. All these limited data suggests that with the effects of colder surface water entering the Mediterranean during H2, cold and arid climate stemming from both the western Mediterranean and North Atlantic influenced the Eastern Mediterranean and some parts of Anatolia. The palaeoclimatic data obtained from different regions and different proxy records confirms that the (sudden and strong) cold and arid conditions identified from the pollen records of northern and southern Anatolia between 25 and 23.5 cal ka BP were related to H2. Thus, the H2 event may have affected some parts of the Anatolian peninsula, as reflected in pollen and lake records. In addition to this, the emergence of the LGM positions of valley glaciers in high mountains in Anatolia after 21.7 ka (except for Dedegöl Mountain, at 24.3 ka BP) (e.g. Zahno et al., 2010) can be related to the effect of H2.

5.2. Environmental conditions between 25 and 23 cal ka BP

5.3. Environmental conditions between 23 and 19 cal ka BP

In this period, the cold and arid climatic conditions in some parts of Anatolia inferred from pollen records showed a steady decrease in AP and a notable increase in NAP. This correlates very well with the timing of H2 event between 25 and 23 cal ka BP (Bard et al., 2000). During H2, a sudden and strong decrease occurred in sea surface temperature (SST) as well as in the salinity of the North Atlantic (Elliot et al., 1998; Bard et al., 2000; Grousset et al., 2000; Voelker et al., 2006) and in the western Mediterranean (Kallel et al., 1997; Paterne et al., 1999; Cacho et al., 2000; Sierro et al., 2005; Melki et al., 2009). The atmospheric conditions over the Mediterranean Sea were affected with dry and cold winds during H2 (Cacho et al., 2000, 2006). Pollen records for the western Mediterranean region show an increase in aridity (Allen et al., 1999; Combourieu-Nebout et al., 2002; Sánchez-Goni et al., 2002). Deep sea drilling cores near southern Iberia revealed the expansion in the semi-desert steppes throughout H2 (Boessenkool et al., 2001; Combourieu-Nebout et al., 2002; Turon et al., 2003; Bout-Roumazeilles et al., 2007). Southward migration of polar air masses over the European continent caused strong cooling episodes at the time of H2, triggering cool glacial conditions there (Guiot et al., 1993; Bigg, 1994; Melki et al., 2009). The Mediterranean Sea surface waters underwent an abrupt decrease of SST at the time of the H2 (Melki et al., 2009). Cold-water input to the Mediterranean originating in the collapse of the North Atlantic Deep Water circulation caused the reduction of evaporation and less precipitation in the Eastern Mediterranean (Bartov et al., 2003). According to Kwiecien et al. (2009), the temperature gradient between mid- and low latitudes stayed high and the westerly jet stayed south, cooler Mediterranean SSTs occurring during Heinrich events would have decreased the atmosphere-sea surface thermal gradient, resulting in a weakening of the Mediterranean low-pressure systems. Subsequently, less frequent and less intense storms and decreased air temperatures would have reduced significantly precipitation in the eastern Mediterranean/Black Sea region. Therefore, the precipitation decrease in the Black Sea region interpreted during H2 can be directly linked to cooling of the Mediterranean (Kwiecien et al., 2009). The effects of H2 seem to have been reflected simultaneously in the lakes in Levant and partly in Anatolia. Maximum levels of Lake Lisan for the Last Glacial were attained at about 25 cal ka BP (Bartov et al., 2002; Landmann et al., 2002). Robinson et al. (2006) proposed that the mean low lake levels of Levant region at about 24 cal ka BP (Landmann et al., 2002; Bartov et al., 2003) is almost

According to pollen records, plant species that indicate cold, cool and humid climate conditions became dominant in some parts of Anatolia (no data for central Anatolia) during the LGM. Despite the dense forest cover that originated in the Marmara, Black Sea and Mediterranean regions between 23 and 19 cal ka BP (AP w% 90e50) pollen data owners (Niklewski and Van Zeist, 1970; Van Zeist et al., 1975; Arslanov et al., 2007) suggests that the climate was cold and arid at that time. Similarly, Van Zeist et al. (1975)
ı as indiinterpreted the pollen records from Karamık Bataklıg cating the dominance of arid and cold conditions in the region throughout this period. However, the fact that the AP percentage reached 30% between 23 and 19 cal ka BP, consisting of Pinus, Alnus, Betula and Abies that thrive in humid conditions, shows an increase in humidity and temperature in the transition zone. In addition, PFT
üt and Ghab indicate that climatic conditions records from Lake Sög were generally cold and humid (AP w50e70%) in this period. The timing of glacier advances observed in the high mountains of Anatolia is consistent with these results. The LGM glaciations occurred between 24.3  1.8 and >17.7  1.4 ka in the Dedegöl Mountains (SW Anatolia) (Zahno et al., 2009) and between 21.5  1.6 and >15.6  1.2 ka in the Kavron Valley and 21.7  1.6 to >16.0  1.2 ka in the Verçenik Valley of the Kaçkar Mountains (NE Anatolia) (Akçar et al., 2007, 2008; parameters recalculated by; Zahno et al., 2009, 2010). However, the onset of deglaciation remains elusive (Zahno et al., 2010). Studies on the Sandıras,
Mountains indicate the beginning of deglaciErciyes, and Uludag ation no later than 20.4  1.3 ka, 21.3  0.9 ka (Sarıkaya et al., 2008, 2009) and 20.3  1.5 ka (Zahno et al., 2010), respectively. Zahno et al. (2010) also propose that all available data related to the last maximum extent of glaciers in Anatolia indicate maximum positions between 24.3  1.8 and 20.3  1.5 ka and coincides with the global LGM interval of 21  2 ka (Mix et al., 2001). This indicates that glacier advancement in the Anatolian mountains in the LGM mostly started after H2. It is known that Anatolia and the surrounding areas were colder than today during the LGM. Hayes et al. (2005) showed that annual SSTs in the Mediterranean and Aegean Sea shores of Anatolia were lower than today (2  C and 2e4  C, respectively). Robinson et al. (2006) estimated that temperature in Anatolia were w8  C less than today. Sarıkaya et al. (2008, 2009) conducted two studies on Erciyes mountain (in Central Anatolia) and Sandıras mountain (in southwestern Anatolia) to identify palaeo-precipitation conditions. According to these studies, the temperature must have been 8e11  C, and 8.5e11.5  C colder than today respectively. These

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values are consistent with the 10  C lower summer temperature recorded by planktonic foraminiferal oxygen isotope values in the Marmara Sea (Mudie et al., 2007). In vegetation models in Europe, AfricaeArabia, and the Middle East and Russian Federation (COHMAP, 1988; Bennett et al., 1991; Peyron et al., 1998; Elenga et al., 2000; Wu et al., 2007), it is estimated that a minimal decrease occurred in the annual precipitation of Anatolia and a cooling of 7e10  C in the winter and of 5  C in the summer occurred. In addition, the mean temperature of the coldest month in this period was generally accepted to be between
14  C and
4  C (in Western and Northern Europe, between
40 and
20  C) (Peyron et al., 1998; Willis and Andel, 2004; Leroy and Arpe, 2007). Therefore, Anatolia represented a refuge area for species, such as Quercus, Castanea sativa and Pinus, which could not survive in Central and Northern Europe during the LGM (Brewer et al., 2002; Taberlet and Cheddadi, 2002; Krebs et al., 2004; Cheddadi et al., 2006). Similarly, Cordova et al. (2009) suggest that some forest was present around the southern and eastern part of the Black Sea. There is no consensus on the humidity levels in Anatolia during
an, 2010). It has been sugthe LGM (e.g. Sarıkaya et al., 2009; Dog gested that the eastern Mediterranean was more arid during the LGM than today through the study of various palaeoclimatic records (e.g. Bar-Matthews and Ayalon, 1997; Robinson et al., 2006) and that Anatolia was more arid than today by using pollen records from plateau regions south of the North Anatolian Mountains (Van Zeist et al., 1975; Van Zeist and Bottema, 1991; Fontugne et al., 1999). Jones et al. (2007) modelled the records obtained from Lake Eski Acıgöl in the Central Anatolia and concluded that precipitation (0.20  0.07 m y
1) during the Glacial (w23e16 14C ka BP) was lower than today (w0.36 m y
1). During the LGM, lake levels reached highstands in the Levant region compared to H2 (Robinson et al., 2006). On the other hand, palaeo-lakes of Anatolia reached high levels during the LGM
lu and Roberts, 1998). Palaeo-Lake Konya reached its (Kuzucuog high level (ca. 20e30 m) between w23.2 and 22.8 cal ka BP (Fontugne et al., 1999). According to Fontugne et al. (1999), there is no material dated between 22.8 and 22 cal ka BP in the Konya Plain, but this period may correspond to a drop in lake level. They reported the highest level of the Palaeo-Lake Konya (20e30 m depth), as indicated by the best preserved beaches, ca. w22 to 20.2 cal ka BP. Around 20.2 cal ka BP, the Palaeo-Lake Konya disappeared completely, probably abruptly (Fontugne et al., 1999). Lake Van level also reached a highstand between 21 and 20 cal ka
lu et al., 2010). BP during the LGM (Kuzucuog Robinson et al. (2006) modelled the precipitation and temperature for Levant region during the LGM. They estimate that temperatures and evaporation were much lower than present, but evaporation was less than precipitation. Therefore, they think that the lower temperatures help to explain how lake levels (e.g. Lake Lisan) could have grown during the glacial phase. Lake highstands (especially Palaeo-Lake Konya highstand) during the LGM in Anatolia were attributed to lowered evaporation rates and the increased catchment runoff resulting from snow-glacier melting
lu and Roberts, 1998; Fontugne et al., 1999; Roberts et al., (Kuzucuog 1999). Although the pollen records reveal that there occur a cold and arid climate during the LGM, the level of Lake Van reached to a highstand supports this view for the Eastern Anatolia. On the other hand, Kashima (2002) suggests that high level of Lake Tuz can be related to increased precipitation as well as lowered evaporation rates due to colder climatic conditions. Similarly, Inoue et al. (1998) suggest that the Konya basin was under a cold
an (2010) used the fluvial and wet climate in glacial epochs. Dog records of the Kızılırmak River in Central Anatolia to suggest that precipitation during the LGM must have been higher than (or

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similar) today’s values. Sarıkaya et al. (2009) estimate that for the glaciers on Mount Erciyes (in Central Anatolia) to form during the LGM, the necessary palaeo-precipitation conditions must have been 20% higher (when the temperature is estimated to have been 8  C lower than today) or 25% lower (when the temperature is estimated to have been 11  C lower than today) than current values. Precipitation in Southwest Anatolia during the LGM would have been 1.9 times more than that of today (Sarıkaya et al., 2008). Similarly, Kwiecien et al. (2009) conclude that the climate was humid in Northwest Anatolia during the LGM. As a result, contrary to previous classical views, the limited amount of PFT data and other proxies seems to support that Anatolian climate (except Eastern Anatolia) during the LGM was more humid than (or similar to) today’s conditions, parallel to the conclusion by Prentice et al. (1992b) for the Mediterranean region from Spain to Iran. Thus, the non-quantitative approach PFT method largely rejects Van Zeist and Bottema’s (1991) views that Anatolia was predominantly cold and arid, only the Black Sea coast belt was forested, the Mediterranean coast, the majority of the Aegean and Marmara regions were covered with steppe forests and the remaining parts were covered with steppe and desert steppe during the LGM. 5.4. Environmental conditions between 19 and 15 ca ka BP The increase in the NAP species in pollen records at 18e16 cal ka BP regarding H1 suggests cold and partly arid climate conditions in some part of Anatolia. This finding is also confirmed by the results derived from proxy data. Palynological records from southern Europe also revealed a decrease in arboreal pollen and an increase in steppe pollen types corresponding with H1 (Reille and Debeaulieu, 1988; Rossignol-Strick and Planchais, 1989; Guiter et al., 2003; Naughton et al., 2007). Similarly, Rossignol-Strick (1995) showed that the Younger Dryas was marked in the Mediterranean and Arabian seas by an increase in Chenopodiaceae. Therefore, they suggest that Younger Dryas was dry and cold in the Mediterranean. Palaeoclimate reconstructions generally suggest that temperature cooler than today during the H1 in the eastern Mediterranean (Bartov et al., 2003; Robinson et al., 2006; Gogou et al., 2007; Roberts et al., 2008; Sarıkaya et al., 2008, 2009; Kwiecien et al., 2009). Kwiecien et al. (2009) state that the initial decrease in carbonate concentration at w18 cal ka BP, indicating a decrease in northwest Anatolia precipitation, which was followed by a more pronounced decrease in carbonate content at 16.4 cal ka BP. They also propose that periods of reduced precipitation are well correlated with low SSTs in the Mediterranean related to H1 and H2. Northward retreat of the polar fronts after w16.4 cal ka BP accounted for warming in the Mediterranean region (Kwiecien et al., 2009). SST derived from the Northern Aegean Sea were as low as 14.5  C in the post-glacial period, then witnessed a sharp increase to 23  C, which most likely corresponds to the Bølling/ Allerød interstadial (Gogou et al., 2007). Roberts et al. (2008) noted that during glacial times Mediterranean lakes deposited carbonates isotopically heavier in d18O compared to the Holocene, partly due to source area effects. They concluded that isotopic enrichment was most marked during intervals corresponding to the H1 and Younger Dryas events, confirming that Late Pleistocene cold stages in the North Atlantic region marked by aridity around much of the Mediterranean. For example, the Eski Acıgöl sequence includes a phase of more positive isotopic values corresponding to the H1 and Younger Dryas stadial (Roberts et al., 2008). Around 18.8 and 18.3 cal ka BP, brackish shallow lake deposits formed in the Konya Plain (Fontugne et al., 1999) and Lake Tuz (Kashima, 2002), respectively. However, in

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general during the post-LGM and Late Glacial stade, depression of the Palaeo-Lake Konya was generally dry with the establishment and re-activation of aeolian dune systems (Fontugne et al., 1999;
lu et al., 1999). Similarly, various data sets seem to suggest Kuzucuog the existence of lake low stands centred about 16e17 cal ka BP, synchronous with H1 in the Levant region (Bartov et al., 2003; Robinson et al., 2006). The lake levels in the Levant peaked after H1 at 15 cal ka BP (Bartov et al., 2002). Late Glacial glaciers readvanced no later than 16.1  1.2 ka ago in
Mountains, and the collapse of the glaciers in the Kovuk the Uludag Valley around 15.1  1.1 ka was a consequence of the pronounced climatic amelioration during the transition to the Bølling interstadial 14.69 ka (Zahno et al., 2010). According to Sarıkaya et al. (2008, 2009), the Late Glacial glacier oscillations happened in the Sandıras Mountains by 16.2  0.5 ka and in the Erciyes Mountains by 14.6  1.2 ka, respectively. The analyses of Late Glacial advance in Central Anatolia during this period suggest that the climate was colder by 4.5e6.4  C and 1.5 times wetter than today (Sarıkaya et al., 2009). According to Sarıkaya et al. (2009) and Zahno et al. (2010), the glacier advance in Anatolia during the Late Glacial was coincident with H1. 6. Conclusion Reinterpretation of previously published pollen records by PFT and non-quantitative biomization methods shows that vegetation and climatic conditions of Anatolia during the Last Glacial were different from previously suggested findings, and not uniform enough to be generalized. Prior to w25 cal ka BP and during w23e19 cal ka BP (during the LGM), a dense forest cover under cold or cool and humid climatic conditions existed in the northern parts of Anatolia (AP percentage 90%) and especially on the coastal belts of Anatolia. Similarly, under humid and cold climatic conditions of the Mediterranean phytogeographic regions in southern Anatolia, a forest cover constituting 50e60% of the entire vegetation was identified. During these periods, cold and arid climate conditions were dominant in Central and Northern Europe, as far as the TurkisheBulgarian border (north of the Istranca Mountains in Turkey). These climatic conditions were accompanied by a vegetation pattern where NAP species were dominant. Similarly, Eastern Anatolia and Western Iran were cold and arid during the LGM. The effects of H2 during the w1500 years between w25 and 23.5 cal ka BP before the local LGM may be reflected in the pollen records of western, northern and southern Anatolia. During this event, cold and arid climatic conditions developed steppe vegetation was dominated in these parts. However, the lack of pollen data from Central and Eastern Anatolia, which have significant mountain ranges as barriers between the coastal belts, is a serious deficiency. The same deficiency is also valid for the Lake Urmia (Western Iran) was cold and arid during the given period. During the Late Glacial, the vegetation pattern and climatic conditions in Anatolia varied by regions, as opposed to the holistic climatic change in the previous periods. The sudden decrease in the AP values during the early period of the Late Glacial at 18e16 cal ka BP and the significant increases in the percentage of NAP revealed that cold and partially arid climate conditions prevailed in some parts of Anatolia. The changes were especially clear in Black Sea
üt and Ghab regions, region and the Marmara Sea, Karamık, Sög where Mediterranean climatic conditions were prevalent. This abrupt but short-lived change may be related to H1, which was influential on a regional scale in the western Mediterranean. The climatic conditions in the Bølling/Alløred interstadial in most of Anatolia were suitable for forest vegetation. At 15 cal ka BP, Anatolia experienced an increase in AP values and a decrease in NAP ratio. Additionally, AP taxa characterizing warm and humid

environments started to become the norm. On the other hand, Eastern Anatolia and the Western Iran highlands were cold and arid until the beginning of the Holocene. Probably, the effects of the Siberian High Pressure systems rather than Mediterranean systems must have delayed the adaptation of plant cover to post-LGM climatic change. The results of the present study reject the hypothesis that the climate in Anatolia and adjacent regions was cold and arid during the LGM and that vegetation consisted largely of steppe and desert steppe. Pollen records used in this study are clearly in accordance with other proxy records for environmental and climatic reconstructions and provided further support for the new interpretations. Acknowledgments We would like to thank Warren Eastwood who revised early text of the manuscript and also Mehmet Akif Sarıkaya and Norm Catto who revised the final English text of the manuscript. We are also grateful to two anonymous reviewers for their constructive comments. References Akçar, N., Schlüchter, C., 2005. Paleoglaciations in Anatolia: a schematic review and first results. Eiszeitalter and Gegenwart 55, 102e121. Akçar, N., Yavuz, V., Ivy-Ochs, S., Kubik, P.W., Vardar, M., Schlüchter, C., 2007. Paleoglacial records from Kavron Valley, NE Turkey: field and cosmogenic exposure dating evidence. Quaternary International 164e165, 170e183. Akçar, N., Yavuz, V., Ivy-Ochs, S., Kubik, P.W., Vardar, M., Schlüchter, C., 2008. A case for a downwasting mountain glacier during Termination I, Verçenik valley. Journal of Quaternary Science 23, 273e285. Allen, J.R.M., Brandt, U., Brauer, A., Hubberten, H.-W., Huntley, B., Keller, J., Kraml, M., Mackensen, A., Mingram, J., Negendank, J.F.W., Nowaczyk, N.R., Oberhansli, H., Watts, W.A., Wulf, S., Zolitschka, B., 1999. Rapid environmental changes in southern Europe during the Last Glacial period. Nature 400, 740e743. Arslanov, K.A., Dolukhanov, P.M., Gei, N.A., 2007. Climate, Black Sea levels and human settlements in Caucasus Littoral 50,000e9000 BP. Quaternary International 167e168, 121e127. _ 1992. The Paleogeography of the Near East from Late Pleistocene to Early Atalay, I., _ Holocene and Human Impact. Ege University Publication, Izmir. Atanassova, J., 2005. Palaeoecological setting of the western Black Sea area during the last 15,000 years. The Holocene 15, 576e584.
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