Palynological Research Research Articles

A comparative palynological study on butterfly mud puddling localities and surface forest samples: a case study from northeast India

This paper presents a palynological examination of surface soil samples procured from butterfly mud puddling locations in northeast India and its comparison with forest surface samples from the immediate vicinity. The modern palynodata from different sites reflect a close similarity in the types and frequency of the pollen grains. However, the diversity of pollen taxa in butterfly mud puddle samples was comparatively higher than the forest surface samples. The pollen recovered in both the butterfly mud puddle location and forest surface samples reflects an evergreen forest composed of Dillenia, Elaeocarpus, Litsea, and Syzygium which exactly displays the current vegetation. The presence of cerealia, along with Brassicaceae, and Hibiscus in the butterfly mud puddle samples is suggestive of the human activity in the vicinity. However, the abundance of Lantana and Melastoma pollen in the mud puddle sediments is a strong indication of the conversion of forest vegetation in to open habitat, because these plants are correlated with human occupation. The extra-regional taxa, especially Rhododendron, in the mud puddle sediments are significant and strongly suggestive of butterfly migration from higher altitudes. Evergreen elements, along with ferns and fungal remains, in the palynoassemblage are strongly indicative of high rainfall under warm and humid climatic conditions. Thus, the main aim of this observation was to identify the potential of butterfly mud puddle samples and its comparison with other surface pollen traps in order to strengthen the relationship between modern pollen and vegetation in floristically rich areas of Assam, northeast India. The palynodata from the butterfly mud puddle locations is a useful source of palynological observation which should be considered in future palynological research.

Evolution of the eucalypts – an interpretation from the macrofossil record

Eucalypts have influenced the fire ecology of the Australian landscape more than any other plant group. They are the iconic plant taxon in the Australian vegetation today, but their origin, early evolution and migration remain poorly understood, mostly because of a remarkably sparse and underworked fossil record. However, a recent major macrofossil find in southern South America, coupled with increasing sophistication of molecular phylogenetic and palynological research allow for a more comprehensive summary of the likely early history of this group of genera. It is likely that the origin was close to the Cretaceous–Paleogene boundary, somewhere in the Weddellian Biogeographic Province (which includes southern South America, western Antarctica and south-eastern Australia), in an area with high natural fire frequency. Evidence for the early record of eucalypts in Australia and their eventual spread across the continent, leading to their current dominance of the Australian plant biomass is growing and is consistent with a drying climate and increasing fire frequency following a very wet period during the Paleogene. The causes of the extinction of eucalypts from South America and probably New Zealand are considered, but remain obscure.

Updated site compilation of the Latin American Pollen Database

The updated inventory of the Latin American Pollen Database (LAPD) offers a wide range of new insights. This paper presents a systematic compilation of palynological research in Latin America. A comprehensive inventory of publications in peer-reviewed and grey literature shows a major expansion of studies over the last decades. The inventory includes 1379 cores and sections with paleoecological data and more than 4800 modern samples from throughout the continent. Through the years, pollen datasets extend over increasing spans of time and show improved taxonomic and temporal resolution. Currently, these datasets are from 12 modern biomes and 30 countries, covering an altitudinal range of 0 to 6300m asl. The most densely sampled regions are the Colombian Andes, the southeast coast of Brazil, and Patagonia. Underrepresented biomes are the warm temperate mixed forest (3%), dry forests (3%), and warm temperate rainforest (1%); whereas steppe, tropical rainforest, and cool grass shrublands, such as the páramos, are generally well represented (all >17%). There are 126 records that span the late Pleistocene to the Last Glacial Maximum transition (21,000calyr BP), and >20% of the records cover the Younger Dryas interval and the Pleistocene/Holocene transition. Reanalysis of numerous sites using multiproxy tools emphasize the informative value of this approach in paleoenvironmental reconstruction. We make suggestions for several pollen sites and regions to be visited again; similarly we identify some key research questions that have yet to be answered. The updated LAPD now provides the platform to support an exciting new phase of global palynological research in which multi-site data are being integrated to address current cutting-edge research questions. The LAPD compilation of sites and the literature database will be available through the Neotoma Paleoecology Database website and a new LAPD website by the end of 2015.

Pollen taphonomy at Shanidar Cave (Kurdish Iraq): An initial evaluation

Caves provide important locations for the study of ancient human activity and environment. One important strand of this ancient environmental work is palynology, yet the taphonomy of pollen in caves is locally contingent and often complex. Shanidar Cave in Kurdish Iraq was the site of important Neanderthal finds and early palynological research, but pollen taphonomy in the cave has not been previously studied, so it is difficult to judge what these ancient pollen assemblages might represent. In this paper, we present pollen from a transect of surface samples within the cave and from comparative surface samples from outside the cave. These show that at present there is a reasonably close correspondence between assemblages accumulating within and in the external environs of the cave, and with the local vegetation. This may suggest that stratigraphic samples may also reflect past local vegetation.

The Blake Event recorded near the Eemian type locality – A diachronic onset of the Eemian in Europe

Multidisciplinary analysis including paleomagnetic, sedimentologic, sea-level change, luminescence dating and palynologic research was performed on a 25 m long orientated core taken at Rutten, close to Eemian key localities in the Netherlands. The main goal of our research was to test a possible delayed onset of temperate conditions in this region compared to Southern Europe, occurring within the Last Interglacial. The sediments revealed the presence of the paleomagnetic Blake Event in ca. 10 m of lower-deltaic floodbasin sediments that contain a pollen record covering the Eemian. The position of the Blake Event in relation to the pollen stratigraphy concurs with the earlier studied Neumark Nord 2 site. Paleomagnetic correlation to core MD95-2042 off SW Iberia indicates ca. 5 kyr diachroneity between the pollen-based onset of temperate interglacial conditions between northern and southern Europe. The onset of the Eemian in north-western and central Europe (ca. 121.0 ka) post-dates the Marine Isotope Stage 6/5e transition by ca. 10 kyr. In addition, the Rutten data provide evidence for a relatively long duration of the Blake Event of at least 8 kyr. The late onset of the temperate conditions that define the base of the Eemian, imply that NW Europe with the Eemian type area is not the most suited region to define the beginning of the Last Interglacial and Late Pleistocene for global chronostratigraphic use.

MIKROSKAMIENIAŁOŚCI ORGANICZNE TERENEWU (KAMBR DOLNY) I PÓŹNEGO EDIAKARU (NEOPROTEROZOIK) OKOLIC KRAKOWA

THE TERRENEUVIAN AND LATE EDIACARAN ORGANIC MICROFOSSILS FROM KRAKOW AREA Abstract. This paper presents the characteristics of the oldest groups of organic microfossils that are documented in two new wells (Trojanowice 2 and Cianowice 2) located on the opposite sides of the fault zone Krakow–Lubliniec separating Upper Silesian and Malopolska blocks). The detail palynological research allowed to determine the age of clastic sediments drilled under Jurassic and Devonian. Two characteristic microfossils assemblages are documented: Late Ediacaran dating the age of the Malopolska Block basement and Terreneuvian known from the oldest Cambrian sediments of the Upper Silesian Block. The documented associations of organic microfossils include the oldest representatives of the bacteria, algae, fungi and animals.

Relicts of a peat cover in the Westerkoggepolder (West Friesland, North-Holland, The Netherlands): The genesis of an eluvial clay soil

This paper presents the result of palynological research of peat relicts, found in the Westerkoggepolder (North-Holland, The Netherlands). In general, such relicts of peat in the actual landscape point to an extensive peat cover in the past that disappeared due to land reclamation and agricultural management. Improvement of the drainage in historical time resulted in shrinkage and bio-oxidation of peat. This caused a land surface lowering of at least 3m and a transformation of Histosols in original peat deposits into eluvial clay covers with Mollic Gleysols. Similar processes occur in other regions in the world were people settled in areas with histosols and reclaimed land for food production. The last relicts of the former peat cover are valuable parts of the national soil archives and deserve the status of geoheritage.

Freeze-dried is as good as frozen: Evaluation of differential preservation of pollen grains in stored lake sediments

In Quaternary studies, sediment storage methods (e.g., freezing, freeze-drying, vacuum-drying) vary across paleoecological disciplines. Typically, palynologists store sediment cores in a freezer or refrigerator; however, sediments already collected and stored (freeze-dried) for paleolimnological studies are available and potentially useful for pollen analysis. Here, we investigate the effect of freeze-drying on pollen grain structure compared to pollen recovered from frozen sediments. Pollen was recorded for eight deterioration categories for paired samples of freeze-dried and frozen sediment obtained from a small alpine lake (Sentinel Lake) in Banff National Park, Canada. No statistically significant differences in pollen deterioration were detected between storage methods for identified pollen taxa and groups (Pinus, Picea, Abies, Alnus, non-arboreal, arboreal). There were also no statistically significant differences between freeze-dried and frozen samples and their mean pollen sum, number of indeterminable pollen grains and number of taxa detected per sample. Instead, deterioration of pollen for both storage methods increased with the depth of sample taken from the core. These results demonstrate freeze-drying as a useful method for storing sediments intended for palynological research. Our findings highlight how limnologists and palynologists can collaboratively share freeze-dried archival sediment samples to reduce field costs and strengthen interpretation of data through analysis of multiple proxies.

Plant remains from the Polish Triassic. Present knowledge and future prospects

ABSTRACT The Triassic plant macrofossils of Poland are very poorly known. There are few Triassic rock exposures here, they contain very few plant specimens, there is little scientific interest in the subject, and the rare plant remains found in drill cores are of low stratigraphical significance. The Lower Triassic macroflora is surprisingly poorer taxonomically than coeval European floras, and only single specimens have been found. The flora of the Middle Triassic is even poorer as a result of the Muschelkalk sea transgression. Only the Upper Triassic floras contain many specimens and taxa. The Upper Triassic macrofloras from Polish territory are well known since the early 19th century. Pioneering descriptions of these floras were given by Goeppert and Raciborski. From the Polish Triassic, the seed fern Lepidopteris ottonis (index species for the Rhaetian stage) and Neocalamites lehmannianus (sphenopsid species typical of almost all European Upper Triassic and Lower Jurassic floras) were described for the first time ever. In the 20th century only single specimens were described from outcrops and drill cores. Barbacka revised Lepidopteris ottonis specimens from old collections and described some new material. Palynological research on Triassic strata in Poland intensified from the 1970s on. That work has produced spore-pollen and megaspore zonations for Triassic strata in Poland, but the correlation of the dispersed spores and pollen grains with their parent plants is low. The Polish Triassic flora is comprised of ferns, lycopsids, sphenopsids, cycads, bennettitaleans, ginkgoaleans and conifers. This flora is taxonomically poorer than equally old and geographically close European floras. All available data about Polish Triassic plants fossils are critically summarised in this paper for the first time. The biostratigraphical and lithostratigraphical correlations of Polish Triassic floras with other European Triassic floras are outlined. New macrofloral assemblages for the Lower and Middle Triassic and macrofloral assemblage zones for the Upper Triassic are proposed for Poland. Recent new finds of taxonomically rich, abundant and well-preserved floras accompanying vertebrate remains in Silesia provide an opportunity for comprehensive research on Polish Triassic floras. This should improve our perception of their taxonomy and allow them to be described in evolutionary and palaeoecological contexts

A multiple profile approach to the palynological reconstruction of Norse landscapes in Greenland's Eastern Settlement

Palynological research is increasingly revealing the landscape impacts of Norse colonisation in southern Greenland. Typically, although not exclusively, these studies are from depositional environments with highly localised pollen source areas close to fjord-side centres of medieval power. In contrast, this paper presents data from Vatnahverfi, an inland district of the Eastern Settlement, and explores the emergence of a cultural landscape through three pollen sequences at variable distances from Norse farms. Two are from mires with small pollen source areas close to (<100m) and distant from (≥1500m) probable farming activities. The other provides a more regional signal of vegetation change, albeit one located close to a Norse settlement. Landnám is marked primarily through an increase in microscopic charcoal and the appearance of pollen from Rumex acetosella, although significant differences between profiles are noted. Close to Norse ruins, pollen productivity from grassland communities increases and woodland and scrub representation declines. Further from archaeological remains, palynologically inferred human activity is primarily characterised by decreased productivity, notably declining influx from woodland and scrub species, reflecting grazing herbivores or coppicing. Abandonment of Vatnahverfi is indicated from the late 14th to early 15th century AD.

The history of early cereal cultivation in northernmost Fennoscandia as indicated by palynological research

The age of the introduction of cereal cultivation in northern Europe has long been debated by researchers from many disciplines, in particular archaeology and palaeoecology. Over the past 40 years extensive palynological data have been collected concerning pre-industrial land use in northern Fennoscandia. This paper reviews palynological studies that include records of fossil cereal pollen from northernmost Sweden, Finland and Norway at latitudes north of 63°N. The geographical extent of known early cultivation sites is constantly expanding, with more than 100 records of cereal pollen pre-dating ad 1700. The oldest records of scattered cereal pollen derive from Neolithic times. Periods of continuous cultivation, indicated by cereal pollen recorded recurrently in the sediment profiles, derive from the Late Neolithic and Early Bronze Age. Collectively, the reviewed pollen records indicate that cereal cultivation was first introduced into areas close to the coast and later to the interior, and that it may have been practiced locally long before sedentary settlements based on intensive cultivation were established during medieval times. The data do not indicate a latitudinal spread of cultivation from south to north. However, methodological problems relating to pollen morphology of cereals, site characteristics and lack of connections to archaeologically excavated sites imply that the value of many early cereal pollen finds remains unclear. To increase our understanding of the context in which cereal cultivation was introduced in northernmost Fennoscandia, multidisciplinary studies integrating palaeoecology, archaeology and history are needed.

Análise Palinológica e a Vegetação

The theme and criteria to be considered during the pollen analysis of quaternary sediments and their interpretation was developed. Techniques for collecting and processing samples, the recognition of botanical taxa through the pollen morphology, knowledge of the age of the material, the physiognomy and phytogeography of vegetation types and their interaction, the experiment reported in previous studies, all together are dominant factors in the quaternary palynological research. The selection and application of this knowledge has been discussed in order to favorable and unfavorable aspects according to the material under study.

Semi-automated segmentation of pollen grains in microscopic images: a tool for three imaging modes

Image analysis methods have the potential to increase the accuracy and rates of data collection in palynological research. Automated segmentation of pollen grains is a method that would facilitate image-based palynological analysis by creating large reference image libraries. We developed an executable for the automated segmentation and cropping of pollen grains from microscope images based on pixel intensity values. Our method works with images taken using transmitted-light, widefield-fluorescence, structured illumination (Apotome), and includes a novel approach for cropping the Apotome Z-stack. The system crops pollen grains from sampled fields of view with ~97% recall and ~97% precision for transmitted-light and widefield-fluorescence images, and ~90% recall and ~89% precision for Apotome fluorescence images. Results differed between different imaging wavelengths for fluorescence images, with Apotome images showing the greatest difference between red and green emission wavelengths. Recall in cropping of transmitted-light images was comparable to previous segmentation efforts.

Presence of the protruding oncus is affected by anther dehiscence and acetolysis technique

A protruding oncus is a projection of the intine in the aperture region. The ubiquitous use of acetolysis in palynological research has led to the presence of a protruding oncus being underreported. Controlled experiments with pollen samples collected from undehisced and dehisced anthers demonstrate that the presence of a protruding oncus is affected by the state of the anther at maturity: dehisced or undehisced and by the preparation technique. In investigating the occurrence of onci, particular attention should be paid both to the dehiscence state of the anthers and the effect of the preparation technique on the intine. Although it has been suggested that protruding onci and pollen buds can be distinguished based on three criteria (size, presence of a large vacuole, separation of the protrusion from the grain), most of these distinctions break down when information is included from more recent studies. Additional study of protruding intinous structures may help clarifing the difference between pollen buds and protruding onci.

Vegetation patterns under climate changes in the Eemian and Early Weichselian in Central Europe inferred from a palynological sequence from Ustków (central Poland)

This paper presents a fresh examination of a pollen profile from Ustków (central Poland), which after preliminary results was thought to span a period between the Early Eemian (MIS 5e) and the upper Pleni-Weichselian (MIS 3). The newly obtained results confirmed the age assessment of the bottom series, but revealed a much older age for the uppermost part of the profile as far back as the Rederstall stadial (MIS 5b). Palynological research showed slightly different patterns of vegetation in comparison to other sites located in Central Europe. Among them were an early Ulmus maximum coinciding with the Betula optimum in the Early Eemian, a relatively late optimum of Taxus baccata during the decline of the Middle Eemian, and a distinct division of the older part of the Late Eemian into phases of Abies-Picea and Picea-Pinus forest domination. The Herning stadial (MIS 5d) falls into a typical bipartition reflected by heathland domination during its older part and the prevalence of Juniperus thickets and Artemisia-Poaceae steppe during its younger part, which makes this succession similar to those from northern Germany. The pollen spectra reflecting the Brørup interstadial (MIS 5c) distinctly revealed an intra-Brørup cold oscillation rarely detected in profiles from Central Europe and a period of Larix dominated forests during the latest part of the interstadial (extraordinarily high percentages exceeding 15% of the total pollen sum). Local pollen taxa, together with the lithological composition of deposits, revealed that there was a water body in the area during the periods of the Late Saalian/Early Eemian transition–Middle Eemian (the decline of the Tilia phase), the Herning stadial (MIS 5d), the early Brørup interstadial (MIS 5c), and the Rederstall stadial (MIS 5b). Between these time-intervals a poor and/or rich fen and/or bog functioned.

Human landscapes and climate change during the Holocene

Palaeoecological and archaeobotanical studies of past environments are essential for understanding the kinds of natural settings in which agriculture and prehistoric societies evolved. They increase our knowledge of how human activities have changed environment and vice versa. Human impact on vegetation cover was recognised from the beginning of modern palynological research (Firbas 1937; Iversen 1941). Based on plant communities described from traditional agriculture and the evidence from archaeobotanical records, Behre (1981) identified palynological anthropogenic indicators. They comprise pollen of cultivated plants together with their associated arable weeds and ruderals. These indicators have been widely used and adapted in different European regions (e.g. Rosch 1996; Brun 2011) and the Near East (e.g. Behre 1990). Apart from reconstructing the vegetation dynamics caused by humans, current palynological research attempts to identify the spatial scale and variation of human activities, and their precise duration and location in given landscapes. Recent advances in quantification and numerical analyses of pollen data including climate reconstruction and vegetation/land-cover modelling, which allow for more precise estimates of the opening of woodlands in past times as well as simulating possible scenarios of human impact on the vegetation and landscapes (e.g. Gaillard et al. 2008). With these methodologies, past and future anthropogenic land cover change can be explored (Gaillard et al. 2010). However, until now most applications of these approaches have mainly concentrated on northwestern Europe. Extending them to further regions would be of great interest. Numerical analyses are of increasing importance for estimating human impact on vegetation and to allow disentangling the human and climatically induced vegetation changes in some cases (Feurdean et al. 2010; Bakker et al. 2012). Climate models can be used for the interpretation of human–environment interactions (e.g. Riehl et al. 2009). However, climatic changes of the last millennia are usually too small in amplitude to be detected by most of the quantitative climate reconstructions using pollen data (Kuhl et al. 2010). Also, microand macro-charcoal records in palaeoecological and archaeobotanical studies serving as climate and human indicators are becoming increasingly relevant (Carcaillet et al. 2002). Investigations of charcoal in soil have focussed on the tropics, the boreal zone and around the Mediterranean, however recently the temperate zone has been integrated as well. In addition, on-site wood charcoal analyses can contribute to reconstructing the human impact on vegetation, especially in areas where no suitable pollen archives are available near human occupation sites (Riehl and Marinova 2008). E. Marinova (&) Centre for Archaeological Sciences, University of Leuven, Celestijnenlaan 200E, bus 2408, 3001 Leuven, Belgium e-mail: elena.marinova@bio.kuleuven.be

Is palynology a credible climate proxy in the Subantarctic?

Pollen and spore analysis is the most successfully used palaeobotanical discipline for reconstructing Holocene vegetation and climate history throughout the world. Subantarctic islands are very specific areas. They are located in the circum-Antarctic Southern Ocean in latitudes that are under strong influence of the southern westerly winds, and are characterised by a treeless, phanerogam-poor flora. Palynological research on many of these islands has resulted in diverging conclusions about how to infer climate history from pollen data. In this study we compare pollen data with macrofossil data on the one hand, and the palaeoenvironmental history based on a multiproxy record on the other hand, of two peat sequences from two different subantarctic islands, South Georgia and Île de la Possession (Îles Crozet). We conclude that palynology must be used with caution as a proxy for climate change on these islands, especially when no other proxy data are available. The upland–lowland principle, as it has been applied in pollen studies in the South Indian Ocean islands, results in erroneous conclusions about climate change on Île de la Possession. More palaeoclimatic multiproxy and pollen studies, in combination with pollen–vegetation relationship studies, can, however, contribute to a more reliable model of how to interpret pollen data in the Subantarctic. We want to stress that our conclusions are only based on Holocene records. Consequently, the question remains if palynology can be used as a palaeoclimatic proxy when climatic changes were more pronounced such as during the last glacial–interglacial transition.

Charcoal identification in species-rich biomes: A protocol for Central Africa optimised for the Mayumbe forest

Direct evidence for Central African vegetation history is mostly derived from palynology and palaeolimnology. Although anthracology has proven worthwhile for palaeovegetation reconstructions in temperate regions and South America, charcoal analysis has hardly been applied for Central Africa. Moreover, a transparent charcoal identification procedure using large databases and well defined characters has never been developed. Therefore, we present a Central African charcoal identification protocol within an umbrella database of species names and metadata, compiled from an on-line database of wood-anatomical descriptions (InsideWood), the database of the world's largest reference collection of Central African wood specimens (RMCA, Tervuren, Belgium) and inventory and indicator species lists. The 2909 Central African woody species covered by this database represent a large fraction of the total woody species richness of Central Africa. The database enables a directed search taking into account metadata on (1) anatomical features, (2) availability of thin sections within the reference collection, (3) species distribution and (4) synonymy. The protocol starts with an anatomical query within this database, focussing on genus rather than species level, proceeds with automatic extension and reduction phases of the resulting species list and ends with a comparative microscopic study of wood reference thin sections and charcoal anatomy. In total, 76.2% of the Central African species in the database are taken into consideration, focussing on indicator and inventory species. The protocol has a large geographical applicability, as it can be optimised for every research area within Central Africa. Specifically, the protocol has been optimised for the Mayumbe region and applied to radiocarbon dated (2055–2205 14Cyr BP) charcoal collections from a pedoanthracological excavation. The validity of the protocol has been proven by the mutual consistency of charcoal identification results and the consistency of these identification results with vegetation history based on phytogeographical and palynological research within and around the Mayumbe. As such, anthracology complements palynology and a combination of both can lead to stronger palaeobotanical reconstructions.

SPOROPOLLEN AND ALGAE ASSEMBLAGES IN DEEP-SEA SEDIMENTS IN THE SOUTH CHINA SEA AND PALEOCLIMATE AND PALEOENVIRONMENT RECORDED IN THEM

Palynological research of deep-sea sediments in the South China Sea has been carried out. Based on the sporopollen and algae assemblages of the deep-sea sediments in cores 149 and 323 in the South China Sea,four sporopollen zones of core 149 have been distinguished in ascending order:Zone 1(407~270 cm):Pinus-Quercus(evergreen)-Umbellferae-Gramineae-Pteris;Zone 2(270~105 cm):Dacrydium-Quercus-Cyathea-Pinus-Polypodiaceae;Zone 3(105~30 cm):Pinus-Polypodiaceae-Quercus(evergreen)-Pteridium;Zone 4(30~0 cm):Gleditsia-Cyathea-Pinus-Polypodiaceae(30~0 cm).Core 323 also has 4 assemblages. Comparison of palynological zone,palaeoclimate,stratigraphy and oxygen isotope of cores 149 and 323 in the South China Sea is done. The sporopollen zones reflect four stages of vegetation,climate and paleoenvironment evolution of the South China Sea since 75 000 years ago. Combined with oxygen isotope dating,the sporopollen and algae data can give scientific evidences for stratigraphic division and reconstruction of paleoclimate and paleoenvironment in the South China Sea.

NEW PALYNOLOGICAL DATA ON THE VEGETATION COMPOSITION DURING THE SULA STAGE OF THE EARLY NEOPLEISTOCENE FOREST STEPPE ZONE OF UKRAINE

The author analyses the published and personal evidences from the palynological research of the Sula sediments (Lower Neopleistocene) and vegetation reconstruction for the recent forest-steppe zone of Ukraine. Proceeding from the studying results (5 sections in the recent forest-steppe zone), the features and regional shifts in the vegetation cover of that time have been recognized for the central parts of the Dnieper-Donets Depression, Ukrainian Shield and Volyn-Podillia Plate. It is proved that the forest-steppe zone existed in the Sula time within the Platform Ukraine as well as the refugia presence near water bodies in the NW and, in part, central regions of Ukraine (thermophyle and broad-leaf plants).