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Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence Lyudmila G. Bezusko a , Sergei L. Mosyakin a, * , Alla G. Bezusko b a M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, 2 Tereshchenkivska Street, Kyiv (Kiev) 01601, Ukraine b National University ‘‘Kyiv-Mohyla Academy’’, 2 H. Skovorody Street, Kyiv (Kiev) 04070, Ukraine article info Article history: Available online 26 July 2008 abstract Pollen assemblages were obtained from deposits from archaeological layers of Early Medieval sites dated to the 3rd–7th and 13th centuries A.D. in the Ovruch Ridge area (Zhytomyr Region, the forest zone of Northern Ukraine). Taxonomic composition of the fossil pollen flora (more than 140 taxa of various ranks) was determined for a narrow temporal interval of the 13th century A.D. Based on the combined evidence of pollen and macrofossils, species composition of the cultivated (13 species) and weedy (38 species) components of the flora for the 13th century was identified. Pollen data are indicative of large-scale deforestation in the Ovruch Ridge area during the 13th century. Ó 2008 Elsevier Ltd and INQUA. 1. Introduction The problem of interaction of humans with their natural envi- ronment in the past may be best approached by combining the methods of natural and historical sciences. On the other hand, reliability of paleobotanical (or, rather, archaeobotanical) recon- structions is much enhanced by combining the evidence of paly- nology and palaeoethnobotany; in other words, by studying microfossils jointly with macrofossils. The use of pollen and spore analysis for the studies of cultural layers of archeological sites, especially in urban areas, provides an insight into the anthropic, or human-influenced, components of past vegetation. This is espe- cially the case when the pollen belongs to plants that are consid- ered as indicators of human economic activities, namely, crops and weeds, as well as other plants indicative of human impact on the environment. Archeological and palynological studies conducted in the southern part of Zhytomyr Polissya (the forest zone of Northern Ukraine), including the Ovruch Ridge, started in 1990s. These studies have determined the first paleopalynological characteristics for cultural deposits of the Early Slavonic settlement of Teterivka dated to the 3rd–7th centuries A.D. (i.e. the second half of the Late Holocene). These data enabled identification of a considerable anthropic impact upon natural vegetation in the area of Teterivka- III settlement during the Early Slavonic period (Bezusko et al., 1994). Palynological studies of the Early Medieval settlements of Ovruch and Gorodets (Bezusko, 2000; Bezusko et al., 2002) also indicate considerable human-induced vegetation changes in the Ovruch Ridge area during the second half of the Late Holocene (SA-3). Zhytomyr Polissya was a key region for the inception and early development of the Ukrainian paleopalynological school founded by Dmytro (Dmitriy) K. Zerov (20.09.1985–20.12.1971). Zerov star- ted his seminal investigations of Holocene deposits in 1930s in the northwestern part of Zhytomyr Oblast’ (administrative region) within its pre-war (WWII) administrative borders (Bezusko, 2003). Palynological evidence obtained from the studies of Holocene deposits in both the Ovruch Ridge and Zhytomyr Polissya is generally in agreement with the main results of the pollen and spore analyses performed in Ukraine in the 20th century. From the viewpoint of conventional (stratigraphy-based) chronology (Zerov, 1934, 1938, 1950; Zerov and Artyushenko, 1961, 1966; Pashkevich, 1963), the level of our present knowledge can be considered as satisfactory. Starting with the 1980s (Chernavskaya and Fogel’, 1989, 1991) palynological studies were increasingly combined with radiocarbon dating. 2. Objectives and methodology The main objective of the study was the reconstruction of vegeta- tion history (both natural and anthropogenic processes) of the Ovruch Ridge (Northern Ukraine) in Early Medieval times. A secondary objective considers the data relative to the participation of Rhododen- dron luteum Sweet in the local vegetation of that time. Pollen analysis was the main method of investigation, considered as the most sensitive instrument for the studies of both natural and human-induced changes in the flora and vegetation of the past. * Corresponding author. E-mail addresses: [email protected] (S.L. Mosyakin), [email protected] (A.G. Bezusko). Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint 1040-6182/$ – see front matter Ó 2008 Elsevier Ltd and INQUA. doi:10.1016/j.quaint.2008.04.016 Quaternary International 203 (2009) 120–128
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Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence

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Page 1: Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence

lable at ScienceDirect

Quaternary International 203 (2009) 120–128

Contents lists avai

Quaternary International

journal homepage: www.elsevier .com/locate/quaint

Flora and vegetation of the Ovruch Ridge (Northern Ukraine)in the Early Middle Ages according to palynological evidence

Lyudmila G. Bezusko a, Sergei L. Mosyakin a,*, Alla G. Bezusko b

a M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, 2 Tereshchenkivska Street, Kyiv (Kiev) 01601, Ukraineb National University ‘‘Kyiv-Mohyla Academy’’, 2 H. Skovorody Street, Kyiv (Kiev) 04070, Ukraine

a r t i c l e i n f o

Article history:Available online 26 July 2008

* Corresponding author.E-mail addresses: [email protected] (S.L. Mosy

(A.G. Bezusko).

1040-6182/$ – see front matter � 2008 Elsevier Ltd adoi:10.1016/j.quaint.2008.04.016

a b s t r a c t

Pollen assemblages were obtained from deposits from archaeological layers of Early Medieval sites dated tothe 3rd–7th and 13th centuries A.D. in the Ovruch Ridge area (Zhytomyr Region, the forest zone of NorthernUkraine). Taxonomic composition of the fossil pollen flora (more than 140 taxa of various ranks) wasdetermined for a narrow temporal interval of the 13th century A.D. Based on the combined evidence ofpollen and macrofossils, species composition of the cultivated (13 species) and weedy (38 species)components of the flora for the 13th century was identified. Pollen data are indicative of large-scaledeforestation in the Ovruch Ridge area during the 13th century.

� 2008 Elsevier Ltd and INQUA.

1. Introduction

The problem of interaction of humans with their natural envi-ronment in the past may be best approached by combining themethods of natural and historical sciences. On the other hand,reliability of paleobotanical (or, rather, archaeobotanical) recon-structions is much enhanced by combining the evidence of paly-nology and palaeoethnobotany; in other words, by studyingmicrofossils jointly with macrofossils. The use of pollen and sporeanalysis for the studies of cultural layers of archeological sites,especially in urban areas, provides an insight into the anthropic, orhuman-influenced, components of past vegetation. This is espe-cially the case when the pollen belongs to plants that are consid-ered as indicators of human economic activities, namely, crops andweeds, as well as other plants indicative of human impact on theenvironment.

Archeological and palynological studies conducted in thesouthern part of Zhytomyr Polissya (the forest zone of NorthernUkraine), including the Ovruch Ridge, started in 1990s. Thesestudies have determined the first paleopalynological characteristicsfor cultural deposits of the Early Slavonic settlement of Teterivkadated to the 3rd–7th centuries A.D. (i.e. the second half of the LateHolocene). These data enabled identification of a considerableanthropic impact upon natural vegetation in the area of Teterivka-III settlement during the Early Slavonic period (Bezusko et al.,1994). Palynological studies of the Early Medieval settlements of

akin), [email protected]

nd INQUA.

Ovruch and Gorodets (Bezusko, 2000; Bezusko et al., 2002) alsoindicate considerable human-induced vegetation changes in theOvruch Ridge area during the second half of the Late Holocene(SA-3).

Zhytomyr Polissya was a key region for the inception and earlydevelopment of the Ukrainian paleopalynological school foundedby Dmytro (Dmitriy) K. Zerov (20.09.1985–20.12.1971). Zerov star-ted his seminal investigations of Holocene deposits in 1930s in thenorthwestern part of Zhytomyr Oblast’ (administrative region)within its pre-war (WWII) administrative borders (Bezusko, 2003).

Palynological evidence obtained from the studies of Holocenedeposits in both the Ovruch Ridge and Zhytomyr Polissya isgenerally in agreement with the main results of the pollen andspore analyses performed in Ukraine in the 20th century. From theviewpoint of conventional (stratigraphy-based) chronology (Zerov,1934, 1938, 1950; Zerov and Artyushenko, 1961, 1966; Pashkevich,1963), the level of our present knowledge can be considered assatisfactory. Starting with the 1980s (Chernavskaya and Fogel’,1989, 1991) palynological studies were increasingly combined withradiocarbon dating.

2. Objectives and methodology

The main objective of the study was the reconstruction of vegeta-tion history (both natural and anthropogenic processes) of the OvruchRidge (Northern Ukraine) in Early Medieval times. A secondaryobjective considers the data relative to the participation of Rhododen-dron luteum Sweet in the local vegetation of that time. Pollen analysiswas the main method of investigation, considered as the most sensitiveinstrument for the studies of both natural and human-induced changesin the flora and vegetation of the past.

Page 2: Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence

Table 1List of Early Medieval sites of the Ovruch Ridge (Northern Ukraine).

No. Site N E Researcher

1. Ovruch 51�190 28�400 T. Bezusko2. Nagoryany-I 51�220 28�310 L. Bezusko3. Gorodets (Gorodische and Gorodky) 51�230 28�140 T. Bezusko4. Pribytky-I 51�170 28�270 L. Bezusko5. Lystvyn 51�200 28�210 L. Bezusko6. Cherevky 51�180 28�240 L. Bezusko7. Norinsk 51�160 28�340 L. Bezusko

Cherevky (№ 2)

Herbs41%

Weeds35%

Cerealia4%

Trees& Shrubs

20%

Fig. 1. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Cherevky No. 2).

Gorodets - Gorodky (№ 8 b)

Trees & Shrubs

87%

Cerealia1%

Herbs4%

Weeds8%

Fig. 2. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Gorodets–Gorodky (No. 8b)).

L.G. Bezusko et al. / Quaternary International 203 (2009) 120–128 121

The palynological studies followed the standard procedure ofpollen preparation and processing proposed by V.P. Grichuk (Pok-rovskaya, 1966). The 15 samples (13 Early Medieval and 2 subfos-sils) were treated with heavy liquids (KIþ CdI2) with specificgravities of 2.0, 2.1, and 2.2. Pollen identifications used the refer-ence pollen collections of the Department of Vascular Plants at theM.G. Kholodny Institute of Botany and of the Department of Biologyat the National University ‘‘Kyiv-Mohyla Academy’’, as well aspollen identification references and special palynomorphologicalpublications (Erdtman, 1943; Monoszon, 1950, 1973, 1976, 1985;Kupriyanova, 1965; Grichuk and Monoszon, 1971; Kupriyanova andAleshina, 1972, 1978; Boros and Jarai-Komlodi, 1975; Bobrov et al.,1983; Artyushenko and Romanova, 1984; Levkovskaya, 1987;Romanova and Bezusko, 1987; Savitsky et al., 1998; Tsymbalyuket al., 2005, 2006). Identification of fossil pollen grains of R. luteumused the results of morphological studies of modern pollensampled in the Ovruch Ridge area (Bezusko et al., 2000). Thebotanical plant names follow the recent nomenclatural checklist ofvascular plants of Ukraine (Mosyakin and Fedoronchuk, 1999), withminor corrections. Pollen percentages were calculated as ArborealPollen (AP)þNon-Arboreal Pollen (NAP)¼ 100% (pollen sum, ortotal pollen). The percentages of spores were calculated from thetotals of pollen and spores taken as 100%. Interpreting the results ofthe paleopalynological analysis of the medieval pollen–spore (PS)samples used methodological principles and data based on theprevious actuopalynological studies (i.e. studies of modern pollen,as opposed to studies of fossil pollen, or paleopalynology). Thesestudies specifically demonstrated the correlation between SPspectra in recent and subrecent surface-soil samples and thepresent-day vegetation patterns in the forest, forest–steppe, andsteppe zones of Ukraine (Zubets’, 1971; Arap, 1972, 1974, 1976;Bezusko et al., 1998, 2004). Arap (1976) conducted methodologicalstudies of this character in Zhytomyr Polissya, while Bezusko et al.(2000) did the same in the Ovruch Ridge area. These studies werecontinued and further developed by the present authors. Thesimilarities established by these studies enabled extrapolation offossil SP data, and reconstruction, with a reasonable degree ofprobability, basic vegetation patterns in the Ovruch Ridge area forthe Early Medieval period.

3. Results

Paleopalynological studies were conducted in the Ovruch Ridgearea (also known as the Slovechno-Ovruch Ridge, Slovechansko-Ovruchskiy Kryazh), which is the most elevated part of ZhytomyrPolissya, reaching 316 m above sea level (Popov, 1968). According tothe geobotanical classification, this area belongs to the ‘CentralPolissya Region’ of the ‘Polissya Subprovince’ of the ‘Central Euro-pean Province of the European Broadleaf Forest Domain’ (Barbar-ych, 1977). Remnants of natural oak-pine and pine forests withR. luteum in the understory still exist in the western part of thisregion (Andrienko, 2006). Palynological characteristics wereobtained for the deposits from cultural layers of Early Medievalsettlements of Ovruch, Gorodets, Lystvyn, Cherevky, Pribytky-I,Nagoryany-I, and Norinsk in the Ovruch Ridge area (all locatedwithin Zhytomyr Region, Ukraine) (Table 1).

According to archeological evidence (Tomashevsky, 1998), EarlyMedieval cultural layers are dated to the 13th century A.D. In the10th–13th centuries A.D. the investigated region was part ofa volost’ (Old Rus’ administrative district) of Vruchii (or Vruchiy,now Ovruch). The medieval Ovruch Princedom was in vassaldependence to the Kievan Rus’ state. Numerous outcrops of rocks,especially pyrophyllite, AlSi2O5(OH), widely used in house building,was among the factors that determined an important role of theOvruch Princedom in the economic life of Kievan Rus’ and neigh-boring states.

The palynological data provided new information on the vege-tation and the influence of human activities on the Ovruch Ridgearea in Early Medieval times (Figs. 1–13). For a narrow time interval(13th century A.D.), the taxonomic composition of the collectivefossil palynoflora (more than 140 taxa of various ranks) was iden-tified (Table 2).

The list of Arboreal Pollen includes 42 taxa (3 identified families,17 genera, and 22 identified species). The forests in the areas of theEarly Medieval settlements in the Ovruch Ridge area were formedmainly by Pinus sylvestris, Quercus robur, Quercus petraea, Carpinusbetulus, Betula pendula, Betula pubescens, and Alnus glutinosa. Lessabundant components of forest communities were Tilia cordata,Acer platanoides, Fagus sylvatica, Fraxinus excelsior, Ulmus cf. sub-erosa, Malus sp., Sorbus sp., Picea abies, and others. Shrubby speciesforming the shrub vegetation layer were Corylus avellana, Sambucusnigra, Viburnum spp., Euonymus sp., Ribes sp., Berberis sp. (probably

Page 3: Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence

Gorodets - Gorodische (№ 8 a)

Herbs20%

Weeds45%

Cerealia7%

Trees & Shrubs

28%

Fig. 3. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Gorodets–Gorodische (No. 8a)).

Lystvyn

Cerealia5%

Weeds5%

Herbs30%

Trees& Shrubs

60%

Fig. 4. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Lystvyn).

Norinsk (№ 1)

Herbs36%

Weeds40%

Cerealia5%

Trees &Shrubs

19%

Fig. 5. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Norinsk No. 1).

Norinsk (№ 2)

Herbs29%

Weeds42%

Cerealia3%

Trees & Shrubs

26%

Fig. 6. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Norinsk No. 2).

Ovruch (№ 2b)

Herbs20%

Weeds43%

Cerealia2%

Trees &Shrubs

35%

Fig. 7. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Ovruch No. 2b).

Pribytky- I (№ 1)

Herbs36%

Weeds40%

Cerealia5%

Trees & Shrubs

19%

Fig. 8. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Pribytky-I (No. 1)).

L.G. Bezusko et al. / Quaternary International 203 (2009) 120–128122

Berberis vulgaris L.), and others. Such species as Salix spp., Callunavulgaris, Ledum palustre, R. luteum, Rubus spp., Humulus lupulus alsoparticipated in formation of the Ovruch Ridge vegetation in the13th century. These species are still typical of modern-day forests inthe forest zone of the central northern part of Ukraine.

The cumulative list of herbaceous plants identified in the SPspectra includes 87 taxa (34 families, 8 genera, and 45 species). Thelist of spores includes 15 taxa (4 identified orders, 4 genera, and 7identified species). The data indicate that in plant communities ofthe Ovruch Ridge in Early Medieval times also participated

cryptogamic vascular plants Lycopodium annotinum, Lycopodiellainundata, Huperzia selago and Botrychium cf. lunaria. Today, thesespecies in the studied area occur rarely and are listed in the RedData Book of Ukraine as rare or endangered taxa (Shelyag-Sosonko,1996).

Results of the paleoethnobotanical analysis of fossil plantremnants from Early Medieval deposits of Gorodets, Lystvyn,Ovruch and Norinsk sites, which had been performed byG.A. Pashkevich were summarized. Species-level pollen identifi-cation enabled combination of the paleopalynological and

Page 4: Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence

Pribytky - I (№ 3)

Herbs30%

Weeds47%

Cerealia8%

Trees&Shrubs15%

Fig. 9. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Pribytky-I (No. 3)).

Pribytky - I (№ 6)

Herbs22%

Trees &Shrubs

30%

Cerealia2%

Weeds46%

Fig. 10. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Pribytky-I (No. 6)).

L.G. Bezusko et al. / Quaternary International 203 (2009) 120–128 123

paleoethnobotanical characteristics for Early Medieval deposits(13th century A.D) of the Ovruch Ridge (Table 3).

4. Discussion

The data provide an insight into the composition of cultivated(13 species) and weedy flora (38 species) in the Ovruch Ridge areaduring the 13th century. Analyses of the herbaceous componentson the species level identify a significant number of weedy plants,making up almost 84% of the total number.

Pribytky -

Herbs30%

Fig. 11. Ratio of pollen in pollen spectrum from cultural la

The data quoted here not only provide the general composition ofthe crop and weed floras in the Ovruch Ridge area in Early Medievaltimes (13th century A.D.) but also clearly demonstrate theadvantage of combining the paleoethnobotanical and palynologicalmethods. The paleoethnobotanical method provides the specificidentification of grain crops (cereals), while palynology can usuallyidentify the pollen of grain crops only to the level of the groupcollectively known as Cerealia. However, palynological materials aremore informative when determining the species composition ofweeds. Undoubtedly, the most reliable taxa are those which pres-ence is independently confirmed by the 2 paleobotanical methods.In this study, these are the weedy species Agrostemma githago,Chenopodium album aggregate, Convolvulus arvensis, Polygonumaviculare aggregate, Fallopia convolvulus, and some cultivated plants,e.g. Cannabis sativa. A. githago, C. album aggregate, F. convolvulus,and C. sativa are listed among paleoethnobotanical finds recorded atOld Rus’ sites in Ukraine (Pashkevich, 1991). It should be noted thatG.A. Pashkevich identified among macrofossils such species of thenative flora as S. nigra, C. avellana, and Rubus idaeus. Pollen grains ofS. nigra, C. avellana, Rubus sp. were also recorded in the pollenspectra of that age. This shows that inhabitants of Early Medievalsettlements were engaged in food gathering and thus activelysupplemented their diet by forest wild plant resources.

Walnut (Juglans regia) pollen, a source of protein-rich food andgood-quality timber, was also identified in the SP spectra, thusproviding the first paleobotanical evidence of the presence of thisvaluable cultivated species in the arboreal flora of the Ovruch Ridgein the 13th century A.D. The pollen data also demonstrate theoccurrence of this cultivated plant in the Kiev’s arboreal flora fromthe second half of the 10th century until the 12th century A.D.(Bezusko et al., 2003).

The pollen of herbaceous plants dominated the spectra ofcultural layers of Early Medieval settlements (13th century A.D.) inthe Ovruch Ridge area. Weedy species and Cerealia hold animportant position among the pollen of herbaceous plants. Signif-icantly, high percentages of pollen of trees and shrubs wereregistered only for the pollen spectra from cultural layers of Gor-odets–Gorodky (No. 8b) (Fig. 2) and Lystvyn (Fig. 4) sites. Thehuman impact on the vegetation in the Gorodets settlement area(Gorodets–Gorodky, No. 8b, Fig. 2) is manifested by high rates ofweed pollen. One may conclude that in the case of Gorodky site, thehuman economic activities have not resulted in an acknowl-edgeable decline of forests. The high percentage AP in the pollenspectrum from the Lystvyn cultural layer is probably due toa sampling bias, as the corresponding sample was taken froma hearth (Fig. 4).

The pollen ratios of arboreal (trees and shrubs) and herbaceousplants in 11 pollen and spore (PS) spectra from subfossil (recent)

I (№ 7)

Weeds49%

Cerealia4%

Trees &Shrubs

17%

yers of Early Medieval settlement (Pribytky-I (No. 7)).

Page 5: Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence

Nagoryany - I (№ 1)

Weeds55%

Herbs22%

Cerealia1%

Trees &Shrubs

22%

Fig. 12. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Nagoryany-I (No. 1)).

Table 2List of microfossils: taxa identified in Early Medieval deposits of the Ovruch Ridgearea (13th century A.D.).

No Taxa (alphabetically by family,and genera within family)

Major synonyms and comments

Pollen grains of trees and shrubsACERACEAE sensu stricto Aceraceae is treated here in the

traditional circumscription. Innew systems it is oftenincluded in Sapindaceae

1. Acer sp.2. Acer platanoides L.

BERBERIDCAEAE3. Berberis sp.

BETULACEAE Including Alnaceae4. Alnus sp.5. Alnus glutinosa (L.) Gaertn.6. Alnus incana (L.) Moench7. Betula sp.8. Betula humilis Shrank9. Betula pendula Roth10. Betula pubescens Ehrn.

CANNABACEAE11. Humulus lupulus L. Humulus lupulus is listed here

because of its liana habit andoriginal association withvegetation of forest margins

CAPRIFOLIACEAE sensu lato Including Viburnaceae andSambucaceae. In new systemsbased on molecular phylogenySambucus and Viburnum aresometimes included inAdoxaceae

12. Caprifoliaceae pollen Genus not identified13. Sambucus sp.14. Sambucus nigra L.15. Viburnum sp.

CELASTRACEAE16. Euonymus sp.

CORYLACEAE Including Carpinaceae17. Carpinus betulus L.18. Corylus avellana L.

ERICACEAE sensu stricto Excluding Pyrolaceae19. Ericaceae pollen Genus not identified20 Calluna vulgaris (L.) Hull21. Ledum palustre L.22. Rhododendron luteum

Sweet(¼Azalea pontica L.)

FAGACEAE23. Fagus sylvatica L.24. Quercus sp.25. Quercus petraea L. ex

Liebl.26. Quercus robur L.

L.G. Bezusko et al. / Quaternary International 203 (2009) 120–128124

soil samples of Zhytomyr Polissya (Arap, 1976) and 2 PS spectra ofsubfossil (recent) soil samples of Early Medieval settlements of theOvruch Ridge (Fig. 14a) were analysed. Results show that subfossilPS spectra 1–9, 12, and 13 belong to the forest type of pollenspectra. Composition of subfossil PS spectra 10 and 11 reflects thechanges in the plant cover under the human influence, because thesamples for methodological studies were taken in treeless areas.The subfossil PS spectrum 12 from the area of Nagoryany-I belongsto the forest type of pollen spectra. However, pollen grains of weedswere also present in that spectrum. In subfossil PS spectrum 13from the area of Cherevky we observed growing quantities ofpollen of herbaceous plants, which was probably caused byanthropic influence. It is confirmed by the presence of abundantpollen of weeds among pollen grains of various herbaceous plants.

Pollen of trees dominate in the PS spectra from deposits of thesecond half of the Late Holocene (SA-3) (Pashkevich, 1963) locatedin the Ovruch Ridge area (Slovechanskoye, Goreloye) and adjacentareas within the 50-km radius (Ozeryanskoye, Nebuga, Wuchky,Poyaskovskoye, Strachov, Ozero Dykoye, Buchmany, KlochkovskoyeGalo, Mostvyanskoye). Subfossil PSs belong to the forest type of PSspectra (Fig. 14b). Pollen grains of herbaceous plants, with consid-erable participation of weedy species, prevail in PS spectra fromcultural layers of Early Medieval settlements (Fig. 14c).

The pollen grains of R. luteum were identified among Ericaceaepollen in PS spectra from cultural layers of Early Medieval settle-ments in the Ovruch Ridge area. Identification used both thestandard pollen reference manuals and the results of special paly-nomorphological studies of modern R. luteum plants sampled in thesame area (Pinetum–Rhododendrosum–Polytrichosum association)

Nagoryany - I (№ 2)

Herbs66%

Weeds20%

Trees &Shrubs

14%

Fig. 13. Ratio of pollen in pollen spectrum from cultural layers of Early Medievalsettlement (Nagoryany-I (No. 2)).

JUGLANDACEAE27. Juglans sp.28. Juglans regia L.

GROSSULARIACEAE29. Grossulariaceae pollen Genus not identified30. Ribes sp.

OLEACEAE31. Fraxinus excelsior L.

PINACEAE32. Picea sp.33. Picea abies (L.) H. Karst.34. Pinus sylvestris L.

ROSACEAE35. Malus sp.36. Rubus sp.37. Sorbus sp.

SALICACEAE38. Salix sp.

Page 6: Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence

Table 2 (continued)

No Taxa (alphabetically by family,and genera within family)

Major synonyms and comments

TILIACEAE In some new systems based onmolecular phylogeny Tiliaceae isincluded in Malvaceae sensu lato

39. Tilia sp.40. Tilia cordata Mill.

ULMACEAE41. Ulmus sp.42. Ulmus cf. suberosa Moench (U. campestris L. aggregate)

Pollen grains of herbsALLIACEAE In some new systems based

on molecular phylogeny Alliaceaeis merged with Amaryllidaceae

43. Alliaceae pollen Genus not identified, mostprobably Allium

APIACEAE44. Apiaceae pollen Genera not identified

ASTERACEAE45. Asteraceae pollen Genera not identified46. Artemisia sp.47. Aster sp.48. Artemisia vulgaris L.49. Centaurea cyanus L.50. Cichorium intybus L.51. Sonchus arvensis L.52. Taraxacum officinale Wigg. aggr.53. Tussilago farfara L.

BORAGINACEAE54. Boraginaceae pollen Genera not identified55. Echium vulgare L.

BRASSICACEAE56. Brassicaceae pollen Genera not identified

CAMPANULACEAE57. Campanulaceae pollen Genus not identified, most

probably Campanula and/orJasione

CANNABACEAE58. Cannabaceae pollen Genus not identified59. Cannabis cf. ruderalis Janisch.60. Cannabis cf. sativa L.

CARYOPHYLLACEAE Including Illecebraceae61. Caryophyllaceae pollen Genera not identified62. Agrostemma githago L.63. Alsine media L. (¼Stellaria media (L.) Vill.)64. Cerastium glomeratum Thuill.65. Dianthus armeria L.66. Dianthus cf. polonicus Zapa1.67. Herniaria polygama J. Gay68. Melandrium cf. album (Mill.)

Garcke(¼Silena alba (Mill.) E. Krause; S.latifolia Poir. subsp. alba (Mill.)Greuter and Burdet)

69. Spergula arvensis L.70. Spergularia rubra (L.) J.

Presl and C. Presl

CHENOPODIACEAE71. Chenopodiaceae pollen Genera not identified72. Atriplex tatarica L.73. Chenopodium album L. aggr.74. Chenopodium cf. foliosum Asch. (¼Blitum virgatum L.)75. Chenopodium hybridum L.76. Chenopodium polyspermum L.77. Chenopodium rubrum L.78. Chenopodium cf. suecicum J.

Murr79 Chenopodium vulvaria L.80 Dysphania botrys (L.) Mosyakin

and Clemants(¼Chenopodium botrys L.)

81. Polycnemum arvense L.82. Kochia laniflora (S. G. Gmel.)

Borbas(¼Kochia arenaria (Maerkl.)Schrad.)

Table 2 (continued)

No Taxa (alphabetically by family,and genera within family)

Major synonyms and comments

CONVOLVULACEAE83. Convolvulus arvensis L.

CRASSULACEAE84. Crassulaceae pollen Genus not identified

CYPERACEAE85. Cyperaceae pollen Genera not identified

EUPHORBIACEAE86. Euphorbiaceae pollen Genus not identified

FABACEAE87. Fabaceae pollen Genera not identified

HALORAGACEAE88. Haloragaceae pollen Genus not identified89. Myriophyllum sp.

HYPERICACEAE90. Hypericaceae pollen Genus not identified, most

probably Hypericum91. Hypericum perforatum L.

LAMIACEAE92. Lamiaceae pollen Genera not identified93. Origanum vulgare L.

LILIACEAE94. Liliaceae pollen Genera not identified

LINACEAE95. Linacea pollen Genus not identified

ONAGRACEAE96. Onagraceae pollen Genera not identified

PAPAVERACEAE97. Papaveraceae pollen Genera not identified98. Chelidonium majus L.

PLANTAGINACEAE sensu stricto99. Plantago sp.100. Plantago lanceolata L.101. Plantago major L.102. Plantago media L.

POACEAE103. Poaceae pollen Genera not identified

Cerealia pollen Collective pollen groupincluding crop cereals (suchas Triticum, Secale, Hordeum,Avena and some others)

POLEMONIACEAE104. Polemoniaceae pollen Genus not identified, most

probably Polemonium caeruleum L.

POLYGALACEAE105. Polygalaceae pollen Genus not identified, most

probably Polygala

POLYGONACEAE106. Polygonaceae pollen Genera not identified107. Fallopia convolvulus (L.) A. Love (¼Polygonum convolvulus L.)108. Polygonum aviculare L. aggr.109. Rumex sp.110. Rumex acetosella L. (¼Acetosella vulgaris Fourr.)111. Rumex confertus Willd.

POTAMOGETONACEAE112. Potamogetonaceae pollen Genus not identified, most

probably Potamogeton

PRIMULACEAE Including Myrsinaceae113. Primulaceae pollen Genera not identified

RANUNCULACEAE114. Ranunculaceae pollen Genera not identified115. Caltha palustris L.116. Ranunculus acris L. aggr.117. Thalictrum flavum L.118. Thalictrum lucidum L.

(continued on next page)

L.G. Bezusko et al. / Quaternary International 203 (2009) 120–128 125

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Table 2 (continued)

No Taxa (alphabetically by family,and genera within family)

Major synonyms and comments

119 Thalictrum simplex L.

PYROLACEAE Often treated withinEricaceae sensu lato

120. Pyrolaceae pollen Genus not identified

ROSACEAE121. Rosaceae pollen Genera not identified

RUBIACEAE122. Rubiaceae pollen Genera not identified

SAXIFRAGACEAE123. Saxifragaceae pollen Genera not identified

SCROPHULARIACEAE sensu lato Scrophulariaceae is treated herein the broad traditional sense

124. Scrophulariaceae pollen Genera not identified

TYPHACEAE sensu stricto Excluding Sparganiaceae125. Typha sp.

URTICACEAE126. Urtica sp.

VALERIANACEAE127. Valerianaceae pollen Genus not identified, most

probably Valeriana128. Valeriana sp.

VIOLACEAE129. Violaceae pollen Genus not identified, most

probably ViolaSpores

BRYALES130. Bryales spores Genera not identified

EQUISETALES: Equisetaceae131. Equisetum sp. Including Hippochaete

LYCOPODIALES132. Lycopodiales spores Genera not identified133. Huperzia selago (L.) Bernh.

ex Schrank and Mart.(¼Lycopodium selago L.)

134. Lycopodium sp.135. Lycopodium annotinum L.136. Lycopodium clavatum L.137. Lycopodiella inundata (L.) Holub (¼Lycopodium inundatum L.)

OPHIOGLOSSALES138. Ophioglossales spores Genus not identified139. Botrychium sp.140. Botrychium cf. lunaria (L.) Sw.

POLYPODIALES141. Polypodiales spores Genera not identified142. Athyrium filix-femina (L.) Roth143. Dryopteris filix-mas (L.) Schott

SPHAGNALES144. Sphagnum sp.

Table 3List of macrofossils and microfossils: taxa identified in Early Medieval deposits of theOvruch Ridge area (arranged according to groups – cultivated plants and weeds)

No. Taxon Macrofossils[G.A.Pashkevich]

Microfossils [L.G.Bezusko and T.V.Bezusko]

Cultivated plants1. Avena sativa L. X Pollen grains of

Cerealia2. Hordeum vulgare L. var.coeleste (coeleste group)

X

3. Hordeum vulgare L. X4. Panicum miliaceum L. X5. Secale cereale L. X6. Triticum dicoccum L. X7. Triticum aestivum L. X8. Triticum monococcum L. X9. Cannabis sativa L. X X10. Pisum sativum L. X –11. Lens culinaris Medik. X –12. Linum usitatissium L. X –13. Juglans regia L. – X

Weeds14. Artemisia vulgaris L. – X15. Centaurea cyanus L – X16. Cichorium intybus L. – X17. Sonchus arvensis L. – X18. Taraxacum officinale Wigg. aggr. – X19. Tussilago farfara L. – X20. Echium vulgare L. – X21. Cannabis cf. ruderalis Janisch. – X22. Agrostemma githago L. X X23. Alsine media L. (Stellaria media

(L.) Vill.)X

24. Cerastium cf. glomeratum Thuill. X25. Melandrium album (Mill.) Garcke X26. Spergula cf. arvensis L. X27. Spergularia rubra (L.) J. Presl and

C. PreslX

28. Atriplex tatarica L. X29. Chenopodium album L. aggr. X X30. Chenopodium foliosum Asch. X31. Chenopodium hybridum L. X32. Chenopodium polyspermum L. X33. Chenopodium rubrum L. X34. Chenopodium vulvaria L.35. Chenopodium suecicum J. Murr X36. Dysphania botrys (L.) Mosyakin and

Clemants (Chenopodium botrys L.)X

37. Polycnemum cf. arvense L. X38. Kochia laniflora (S.G. Gmel.) Borbas X39. Convolvulus arvensis L X X40. Chelidonium majus L. X41. Plantago lanceolata L. X42. Plantago major L. X43. Fallopia convolvulus (L.) A. Love X X44. Polygonum aviculare L. aggr. X X45. Rumex acetosella L. X46. Rumex confertus Willd. X47. Bromus secalinus L. X –48. Echinochloa crusgalili (L.) P. Beauv. X49. Setaria glauca (L.) P. Beauv. X –50. Setaria viridis (L.) P. Beauv. X –51. Galium aparine L. X –

L.G. Bezusko et al. / Quaternary International 203 (2009) 120–128126

(Bezusko et al., 2000). The data show that pollen grains of R. luteumparticipated in the PS spectra of cultural layers of Early Medievalsettlements of Gorodets (Gorodky), Cherevky, Nagoryany-I, andOvruch (0.5–4.0% of total pollen grains).

The problem of origin and distribution of R. luteum in UkrainianPolissya (Ukraine’s northern forest zone) remains a debated issue inUkrainian historical plant geography. Most scholars share the viewabout the relict status of this species in Ukrainian Polissya, includingthe Ovruch Ridge. However, their opinions differ regarding the age ofthis presumably relict species, ranging widely from the Tertiary, tothe Riss–Wurm or even the Holocene, or even suggesting its veryrecent introduced status. Opinions also differ as about possiblemigration pathways of this species into Ukrainian Polissya, where itoccurs in isolated localities more than 800 km away from the area ofits continuous range. Paleopalynological data cannot provide

answers to all these complicated questions; hopefully, they will beeventually solved, by means of focused joint paleobotanical andphylogeographical studies. However, the presently available dataprovide evidence in favor of rejection of the hypothesis of the recentintentional introduction of this species into Ukrainian Polissya.Palynological and paleoclimatic data on the periglacial flora andvegetation of Ukraine (Tarasov, 2000; Tarasov et al., 2000) during theLast Glacial Maximum give little ground for the concept of eitherTertiary or Riss–Wurm ages (Barbarych, 1953, 1962; Smyk, 1964,1965; Udra, 1982; Smyk and Bortnyak, 1984) of presumably relictspecies, such as R. luteum, Hedera helix L., and Q. petraea.

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0

20

40

60

80

100

Trees & Shrubs 95 96 94 97 95 98 99 97 93 74 70 94 66Herbs 5 4 6 3 5 2 1 3 7 26 30 6 34

1 2 3 4 5 6 7 8 9 10 11 12 13

0

50

100

Trees & Shrubs 88 96 92 92 85 85 90 95 79 88 93

Herbs 12 4 8 8 15 15 10 5 21 12 7

1 2 3 4 5 6 7 8 9 10 11

0

20

40

60

80

100

Trees & Shrubs 35 14 22 30 87 19 15 30 17 60 20 19 26Herbs 65 86 78 70 13 81 85 70 83 40 80 81 74

1 2 3 4 5 6 7 8 9 10 11 12 13

a

b

c

Fig. 14. Ratios of pollen grains of arboreal and herbaceous plants in pollen and sporespectra from (a) subfossil samples, (b) Holocene deposits (SA-3), and (c) cultural layersof the Ovruch Ridge.

L.G. Bezusko et al. / Quaternary International 203 (2009) 120–128 127

5. Conclusions

1. The pollen data obtained for archaeological deposits at sevenEarly Medieval settlements of the Ovruch Ridge reveal thegeneral pattern of natural and anthropogenic changes in theplant cover within a narrow temporal interval of the 13thcentury A.D.

2. Generalized pollen data (subfossil PS spectra, PS spectra ofHolocene deposits – SA-3, and PS spectra of cultural layers)reveal the general character of the Early Medieval spectra astypical of open (non-forest or even treeless) landscapes. Pollenof weedy species plays an important role in the herbaceouscomponent of PS spectra. Palynological data indicate a large-scale deforestation in the Ovruch Ridge area during the 13thcentury.

3. Taxonomic composition of the collective fossil palynofloras(more than 140 taxa of various ranks) was determined fora narrow temporal interval of the 13th century A.D.) Based onthe combined evidence of palaeopalynological and paleo-ethnobotanical analyses, we identified the species compositionof the cultivated (13 species) and weedy (38 species) floras forthe 13th century.

4. The data obtained for macro- and microfossils demonstrate thehigh potential of combined paleoethnobotanical and palyno-logical approach. The paleoethnobotanical method providesdetailed information regarding crop species. At the same time,

paleopalynological materials are more informative withrespect to the species composition of the weedy flora. The mostreliably identified taxa are those which presence is confirmedby the both paleobotanical methods. In our study, these aresuch weedy species as A githago, C. album aggr., C. arvensis,P. aviculare, F. convolvulus, as well as several cultivated plants,e.g. C. sativa.

5. Pollen of the walnut tree, J. regia, a source of edible nuts andhigh-quality timber, was identified in the PS spectra. Thispalynological data is the first paleobotanical evidence for thepresence of this valuable cultivated species in the Ovruch Ridgearboreal flora already as far back as the 13th century A.D.

6. The pollen records of the Early Medieval deposits of the OvruchRidge indicate the presence of R. luteum, the controversialspecies that had been viewed either as a recent introduction ora relict. Our data do not confirm the opinion about the humanintroduction of this species into the northern forest zone ofUkraine. The age of other presumed relicts in the studied area(H. helix L., Q. petraea L. ex Liebl., and others) should also bereconsidered.

Acknowledgments

The authors express their sincere gratitude to Dr. AndreyP. Tomashevsky for providing samples for our palynological inves-tigations and their archaeological dating, Prof. Galina A. Pashkevichfor her original paleoethnobotanical data on Gorodets, Lystvyn,Ovruch and Norinsk sites, Olga M. Kornienko for her assistance inpreparing the illustrations, and Timur V. Bezusko for providingoriginal palynological data for the Ovruch and Gorodets sites.

References

Andrienko, T.L. (Ed.), 2006. Fitoriznomanittya Ukrayins’kogo Polissya ta yogookhorona (Phytodiversity of Ukrainian Polissya and its Conservation). Phyto-sociocenter, Kiev, p. 316 (in Ukrainian).

Arap, R.Ya., 1972. Palynological investigations of soil surface layers in the forest–steppe zone of the Ukrainian SSR. Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.)29 (4), 506–513 (in Ukrainian).

Arap, R.Ya., 1974. Correlation of recent spore and pollen spectra and the plant covercomposition in Volhynian Polissya. Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.)31 (1), 493–498 (in Ukrainian).

Arap, R.Ya., 1976. Palynological investigations of soil surface layers in UkrainianPolesie. In: Teslenko, I.V. (Ed.), Palinologicheskie issledovaniya osadochnykhotlozheniy Ukrainy I smezhnykh territoriy (Palynological Investigations ofSedimentary Layers of Ukraine and Adjacent Regions). Naukova Dumka, Kiev,pp. 11–16 (in Russian).

Artyushenko, A.T., Romanova, L.S., 1984. Morfologiya pyl’tsy reliktovykh, ende-michnykh i redkikh vidov flory Ukrainy. (Morphology of Pollen of Relict,Endemic and Rare Species of the Flora of Ukraine). Naukova Dumka, Kiev, 48 pp.(in Russian).

Barbarych, A.I., 1953. Distribution of yellow rhododendron in Ukrainian Polissyaand prospects of its economic use. Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.)10 (2), 55–60 (in Ukrainian).

Barbarych, A.I., 1962. Yellow rhododendron, a relict of the Tertiary flora in UkrainianPolissya. Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.) 19 (2), 30–39 (inUkrainian).

Barbarych, A.I. (Ed.), 1977. Geobotanichne rayonuvannya Ukrayins’koyi SSR(Geobotanical Subdivision of the Ukrainian SSR). Naukova Dumka, Kiev, p. 303.

Bezusko, A.G., 2003. Paleobotanical Studies of Holocene Deposits of ZhytomyrPolissya in Works of D.K. Zerov. Naukovi Zapysky NaUKMA. Gumanitarni Nauky.(Sci. Transact. National Univ. Kiev-Mohyla Academy. Humanities), vol. 22, part2. KM Academia, Kiev, pp. 171.C–174.C.

Bezusko, L.G., Bezusko, A.G., Yesilevsky, S.O., 1998. Actuopalynological Aspects ofthe Holocene Paleoecology (on the Example of the Steppe Zone of Ukraine).Naukovi Zapysky NaUKMA. Pryrodnychi Nauky (Sci. Transact. National Univ.Kiev-Mohyla Academy. Natural Sciences), vol. 5, pp. 51–57 (in Ukrainian).

Bezusko, L.G., Bezusko, A.G., Mosyakin, S.L., 2004. Modern Status of PalynologicalStudies of Subfossil Soil Layers of the Steppe Zone of Ukraine. Stepovi i galo-fil’ni ekosystemy Ukrayiny (Steppe and Halophilous Ecosystems of Ukraine. Sci.Collection Devoted to the 100th Anniversary of Prof. G.I. Bilyk). M.G. KholodnyInstitute of Botany, National Academy of Sciences of Ukraine, Kiev, pp.398–414.

Page 9: Flora and vegetation of the Ovruch Ridge (Northern Ukraine) in the Early Middle Ages according to palynological evidence

L.G. Bezusko et al. / Quaternary International 203 (2009) 120–128128

Bezusko, L.G., Bezusko, T.V., Mosyakin, S.L., 2003. A partial reconstruction of theflora and vegetation in the central area of Early Medieval Kiev, Ukraine, basedon the results of palynological investigations. Urban Habitats 1 (1), 15 (SpecialIssue: Urban Floras) Available from: <www.urbanhabitats.org/v01n01/nonnativekiev_pdf.pdf>.

Bezusko, L.G., Bezusko, T.V., Mosyakin, S.L., Tomashevsky, A.P., 2002. Palynologicaland archeological characteristics of deposits in the area of early medievalOvruch (Zhytomyr Region, Ukraine). In: V International Conference onAnthropization and Environment of Rural Settlements. Flora and Vegetation.Abstracts. Uzhgorod and Kostryno, Ukraine, pp. 13–15.

Bezusko, L.G., Tomashevsky, A.P., Pashkevich, G.O., Timchenko, T.V., 1994. Paleo-botanical studies of deposits of the Early Slavonic settlement of Teterivka-III(Zhytomyr Region, Ukraine). Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.) 51 (2/3), 157–164 (in Ukrainian).

Bezusko, T.V., 2000. On Natural Conditions of Medieval Settlements on the OvruchRidge (Palynological Data). Naukovi Zapysky NaUKMA. Spets. Vypusk (Sci.Transact. National Univ. Kiev-Mohyla Academy. Special Issue), vol. 18, pp. 291–293 (in Ukrainian).

Bezusko, T.V., Martynyik, O.O., Popovich, S.Yu., 2000. Some Aspects of the Use ofPollen of Rhododendron luteum Sweet (Ericaceae) in the Flora of Ukraine forSpore and Pollen Analysis. Naukovi Zapysky NaUKMA. Biologiya and Ekologiya(Sci. Transact. National Univ. Kiev-Mohyla Academy. Biology and Ecology), vol.18, pp. 12–17 (in Ukrainian).

Bobrov, A.E., Kupriyanova, L.A., Litvintseva, M.V., Tarasevich, V.F., 1983. Sporypaporotnikoobraznykh i pyl’tsa golosemennykh i odnodol’nykh rasteniy floryevropeyskoy chasti SSSR. (Spores of Ferns and Fern Allies, and Pollen ofGymnosperms and Monocots of the Flora of the European Part of the USSR).Nauka, Leningrad, 208 pp. (in Russian).

Boros, A., Jarai-Komlodi, M., 1975. An Atlas of Recent European Moss Spores. Aka-demiai Kiado, Budapest.

Chernavskaya, M.M., Fogel’, G.A., 1989. Climatic variability in the South-West of theEuropean part of the USSR in the Holocene. Doklady AN SSSR (Transactions ofthe Academy of Sciences of the USSR) 307 (6), 1474–1477.

Chernavskaya, M.M., Fogel’, G.A., 1991. Reconstruction of the climate of Polesie andCis-Carpathian area according to palynological data. Izvestiya AN SSSR. Ser.Geogr. (Proceedings of the Academy of Sciences of the USSR, Ser. Geogr.) 1991(2), 98–105. in Russian.

Erdtman, G., 1943. An Introduction to Pollen Analysis. Chronica Botanica Co., Wal-tham, MA.

Grichuk, V.P., Monoszon, M.Kh., 1971. Opredelitel’ odnolychevykh spor papor-otnikov iz semeystva Polypodiaceae R. Br., proizrastayushchikh na territoriiSSSR. (Identification Manual of Uniseriate Spores of Ferns of the Family Poly-podiaceae R. Br. Occurring in the Territory of the USSR). Nauka, Moscow, 124 pp.(in Russian).

Kupriyanova, L.A., 1965. Palinologiya serezhkotsvetnykh. Palynology of Amentiferae.Nauka, Moscow and Leningrad, 214 pp. (in Russian).

Kupriyanova, L.A., Aleshina, L.A., 1972. Pyl’tsa i spory rasteniy flory Evropeyskoychasti SSSR. (Pollen and Spores of Plants in the Flora of the European Part of theUSSR), vol. 1. Nauka, Leningrad, 170 pp. (in Russian).

Kupriyanova, L.A., Aleshina, L.A., 1978. Pyl’tsa dvudol’nykh rasteniy flory Evro-peyskoy chasti SSSR. Lamiaceae–Zygophyllaceae. (Pollen of DicotyledonousPlants in the Flora of the European Part of the USSR. Lamiaceae–Zygophylla-ceae). Nauka, Leningrad, 183 pp. (in Russian).

Levkovskaya, G.M., 1987. Priroda i chelovek v srednem golotsene Lubanskoy niziny.(Nature and Man in the Middle Holocene of the Lubanskaya Lowland). Zinatne,Riga, 94 pp. (in Russian).

Monoszon, M.Kh., 1950. Description of pollen of Artemisia species growing in theterritory of the USSR (for pollen analysis). In: Trudy Geol. Instituta (Proceedingsof Geological Institute), vol. 46, pp. 271–360 (in Russian).

Monoszon, M.Kh., 1973. Opredelitel’ pyl’tsy vidov semeystva marevykh. (PollenIdentification Manual of Species of the Family Chenopodiaceae). Nauka, Mos-cow, 94 pp. (in Russian).

Monoszon, M.Kh., 1976. Diagnostics of pollen of species of the genus Thalictrum L..In: Palinologiya v SSSR (Palynology in the USSR). Nauka, Moscow, pp. 24–27 (inRussian).

Monoszon, M.Kh., 1985. Pollen morphology of species of the genus Plantago L. (amanual for spore and pollen analysis). In: Volkova, V.S., Khlonova, A.F. (Eds.),Palinostratigrafiya mezozoya i kaynozoya Sibiri (Palynostratigraphy of theMesozoic and Cenozoic of Siberia). Nauka, Novosibirsk, pp. 65–73 (in Russian).

Mosyakin, S.L., Fedoronchuk, M.M., 1999. Vascular Plants of Ukraine. A Nomencla-tural Checklist. National Academy of Sciences of Ukraine, Kiev, xxivþ 346 pp.

Pashkevich, G.O., 1963. History of vegetation of Zhytomyr Polissya in the Holoceneaccording to data of spore–pollen studies. Ukrayinski Botanichnyi Zhurnal (Ukr.Bot. J.) 20 (6), 52–62 (in Ukrainian).

Pashkevich, G.A., 1991. Paleoetnobotanicheskie nakhodki na territorii Ukrainy.Drevnyaya Rus’. Katalog. (Paleoethnobotanical Finds in the Territory of Ukraine:

Ancient Rus’. Catalogue). Institute of Archaeology, National Academy ofSciences of Ukraine, Kiev, 45 pp. (in Russian).

Pokrovskaya, I.M. (Ed.), 1966, Paleopalinologiya (Paleopalynology), vol. 1. Nauka,Leningrad, p. 351 (in Russian).

Popov, V.P. (Ed.), 1968. Fiziko-geograficheskoe rayonirovanie Ukrainskoy SSR(Physiographic Subdivision of the Ukrainian SSR). Kiev University Press, Kiev, p.631 (in Russian).

Romanova, L.G., Bezusko, L.G., 1987. Pollen of some weedy plants of the familyCaryophyllaceae Juss. as an indicator of anthropic activity in the past.Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.) 43 (1), 42–46 (in Ukrainian).

Savitsky, V.D., Bezusko, L.G., Savitska, O.V., Bezusko, T.V., 1998. Pollen of weedyspecies of Papaveraceae Juss. as an indicator of anthropic activity in the past.Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.) 55 (3), 301–306 (in Ukrainian).

Shelyag-Sosonko, Yu.R. (Ed.), 1996. Chervona knyga Ukrayiny. Roslynnyy svit (RedData Book of Ukraine. Plants). Ukraine Encyclopedia Publications, Kiev, p. 604(in Ukrainian).

Smyk, G.K., 1964. Interesting floristic finds in the Slovechno-Ovruch Ridge.Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.) 21 (4), 101–103 (in Ukrainian).

Smyk, G.K., 1965. An overview of the flora of the Slovechno-Ovruch Ridge.Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.) 22 (4), 63–67 (in Ukrainian).

Smyk, G.K., Bortnyak, N.N., 1984. Floristic finds in the Slovechno-Ovruch Ridge(Central Polessie of the Ukrainian SSR). Botanicheskiy Zhurnal (Bot. J., Lenin-grad) 69 (18), 1096–1099 (in Russian).

Tarasov, P.E., 2000. Reconstruction of the climate and vegetation of Northern Eur-asia in the Late Pleistocene according to palynological data. In: Problemypaleogeografii i stratigrafii pleystotsena (Problems of Paleogeography andStratigraphy of the Pleistocene). Moscow University Press, Moscow, pp. 70–96(in Russian).

Tarasov, P.E., Volkova, V.S., Webb III, T., Quiot, J., Andreev, A.A., Bezusko, T.V.,Bezusko, L.G., Bykova, G.V., Dorofeyuk, N.I., Kvavadze, E.V., Osipova, I.M.,Panova, N.K., Sevastyanov, D.V., 2000. Last Glacial Maximum biomes recon-structed from pollen and plant macrofossil data from Northern Eurasia. Journalof Biogeography 27, 609–620.

Tomashevsky, A.P., 1998. Preliminary results of archaeological studies of theOvruch Archaeological Expedition in 1996–1997 within the area of OvruchDistrict, Zhytomyr Region. In: Arkheologichni vidkryttya v Ukrayini 1997–1998rr. (Archaeological Discoveries in Ukraine, 1997–1998). Institute of Arch-aeology, National Academy of Sciences of Ukraine, Kiev, pp. 45–48 (inUkrainian).

Tsymbalyuk, Z.M., Mosyakin, S.L., Bezusko, L.G., 2005. New Approaches to Devel-oping an Identification Manual of Pollen of Chenopodiaceae for Pollen Analysis(Taxa Occurring in Ukraine). Naukovi Zapysky NaUKMA. Biologiya ta Ekologiya(Sci. Transact. National Univ. Kiev-Mohyla Academy. Biology and Ecology), vol.43, pp. 19–25 (in Ukrainian).

Tsymbalyuk, Z.M., Mosyakin, S.L., Bezusko, L.G., 2006. Pollen morphology of thegenus Plantago l. s. l. (Plantaginaceae Juss. s. str.) in the flora of Ukraine forspore–pollen analysis. Naukovi Zapysky NaUKMA. Biologiya ta Ekologiya (Sci.Transact. National Univ. Kiev-Mohyla Academy. Biology and Ecology), vol. 54,pp. 24–30 (in Ukrainian).

Udra, I.F., 1982. Dispersal of trees, their migrational capacity, and biogeographicalinterpretation of Quaternary events. Botanicheskiy Zhurnal (Bot. J., Leningrad)67 (8), 1047–1059. in Russian.

Zerov, D.K., 1934. Time and conditions of development of sphagnum bogs ofnorthwestern Ukraine. Zhurnal Institutu Botaniki VUAN (Journal of theInstitute Botany of the All-Ukrainian Academy of Sciences) 2 (10), 137–153(in Ukrainian).

Zerov, D.K., 1938. Bolota URSR. Roslynnist’ i stratigrafiya. (Mires and Bogs of theUkrainian SSR. Vegetation and Stratigraphy). Academy of Sciences of theUkrainian SSR Publications, Kiev, 164 pp. (in Ukrainian).

Zerov, D.K., 1950. Main features of postglacial history of vegetation in the territoryof the Ukrainian SSR. In: Trudy konferentsii po sporovo-pyl’tsevomu analizu(1948 g.) (Proceedings of the Conference on Spore–Pollen Analysis, 1948).Moscow, pp. 43–61 (in Russian).

Zerov, D.K., Artyushenko, A.T., 1961. History of vegetation of Ukraine since themaximum glaciation according to data of spore–pollen analysis. In: Chetver-tichnyy Period (The Quaternary). Issue 13–15. Academy of Sciences of theUkrainian SSSR, Kiev, pp. 300–322 (in Russian).

Zerov, D.K., Artyushenko, O.T., 1966. Spore–pollen studies of lake and mire depositsof the Ovruch Ridge. In: Paleogeografichni umovy terytoriyi Ukrayiny v pliot-seni ta antropogeni (Paleogeographic Conditions within the Territory of Ukrainein the Pliocene and Anthropogene). Geographic Sector of the Academy ofSciences of the Ukrainian SSR and Geographic Society of the Ukrainian SSR,Kiev, pp. 20–25 (in Ukrainian).

Zubets’, R.Ya., 1971. Spore and pollen studies of subfossil soil layers of the steppepart of Ukraine. Ukrayinski Botanichnyi Zhurnal (Ukr. Bot. J.) 28 (2), 192–198 (inUkrainian).