Arctic plant diversity in the Early Eocene greenhouse

For the majority of the Early Caenozoic, a remarkable expanse of humid, mesothermal to temperate forests spread across Northern Polar regions that now contain specialized plant and animal communities adapted to life in extreme environments. Little is known on the taxonomic diversity of Arctic floras during greenhouse periods of the Caenozoic. We show for the first time that plant richness in the globally warm Early Eocene (approx. 55-52 Myr) in the Canadian High Arctic (76° N) is comparable with that approximately 3500 km further south at mid-latitudes in the US western interior (44-47° N). Arctic Eocene pollen floras are most comparable in richness with today's forests in the southeastern United States, some 5000 km further south of the Arctic. Nearly half of the Eocene, Arctic plant taxa are endemic and the richness of pollen floras implies significant patchiness to the vegetation type and clear regional richness of angiosperms. The reduced latitudinal diversity gradient in Early Eocene North American plant species demonstrates that extreme photoperiod in the Arctic did not limit taxonomic diversity of plants.     

The environmental implications of upper Paleozoic plant-fossil assemblages with mixtures of wetland and drought-tolerant taxa in tropical Pangea

We evaluate the influences of elevation and climate on the spatio-temporal distribution of wetland and dryland biomes during the Pennsylvanian and early Permian in tropical Pangea. The longstanding “upland model” places drought-tolerant vegetation in elevated habitats, where slope and drainage created moisture-limited substrates under a humid climate that simultaneously promoted peat accumulation in contemporaneous lowlands. Upland plants were periodically transported to, and buried in, lowlands. Rare preservation of dryland vegetation thus reflects its general absence in basins, and taphonomic vagaries of long-distance transport. The alternative “climate model” proposes that drought-tolerant plants dominated tropical habitats when climate was seasonally dry, with wetland vegetation reduced to scattered refugia. Environmental changes attending glacial-interglacial cycles caused alternating wetter-drier conditions, and the relative abundance of wetland versus dryland biomes in basinal lowlands thus varied with climatic oscillations. The paucity of drought-tolerant plants reflects a preservational megabias against habitats with seasonal moisture deficits.The environmental signal of “mixed” plant-fossil assemblages, comprising taxa characteristic of both wetland and dryland biomes, may help resolve these debates. We review key Pennsylvanian and lower Permian mixed assemblages from tropical Euramerican Pangea, and interpret their original habitats and climatic contexts based on multidisciplinary lines of evidence, including sedimentology, taphonomy, physiology, and paleoecology. Evaluations also consider patterns of vegetational distribution and taphonomy in modern tropical environments. We suggest that even a cursory view of current tropical plant distribution exposes flaws in the upland model. Where tropical climate is sufficiently humid to support peat swamps, slopes and elevated habitats do not host drought-tolerant vegetation, but are occupied by plants similar to those in lowland settings. This occurs because equable, high precipitation strongly dampens water-table variation across entire landscapes. Furthermore, taphonomic studies indicate that most plant-fossil assemblages record vegetation living near the burial site. Fossil floras thus reflect environmental conditions near their growth site, excluding an upland origin for most occurrences of drought-tolerant taxa. Conversely, the climate model is consistent with modern tropical vegetational distribution and soundly explains late Paleozoic floristic patterns. When Pangean tropical lowlands experienced seasonally dry conditions, plants tolerant of moisture deficits dominated most habitats, whereas wetland vegetation was restricted to wetter sites with greater preservation potential. This occurred because topographic variations are magnified under seasonal precipitation regimes, creating a complex habitat mosaic with wetland patches in a landscape subject to seasonal drought. Accordingly, we propose that a macrofloral assemblage with even rare drought-tolerant taxa indicates seasonality in the broader landscape.At larger spatio-temporal scales, disagreement also persists about whether tectonic uplift or long-term climatic drying was the primary driver of changes in late Paleozoic floristic patterns and areal extent of tropical peat swamps. We argue that tectonic activity alone cannot explain the drastic reduction in peat swamps or coincident changes in dominance-diversity of wetland vegetation. Rates of plant dispersal and evolution far outpace that of mountain building, and peat-forming wetlands persisted in elevated habitats well into the Late Pennsylvanian. Therefore, progressive late Paleozoic aridification was the most probable driver of changing floral patterns and the distribution of wetland and dryland biomes in tropical Pangea.     

Floristic and vegetational changes in the Iberian Peninsula during Jurassic and Cretaceous

The successive vegetations inhabiting the Iberian Peninsula from the Triassic/Jurassic boundary to the Cretaceous/Tertiary Boundary is reviewed based on published palynological and macrofloral data, and the vegetational changes set in a palaeogeographical and climate context. Xerophytic microphyllous coniferous forests and pteridophyte communities of arid environments dominated the Jurassic and earliest Cretaceous vegetation. This vegetation was replaced in the mid Early Cretaceous by mixed forests of pteridophytes, gymnosperms and angiosperms. The composition of the successive plant assemblages suggests that a subtropical arid climate generally prevailed in the Iberian Peninsula during Jurassic-Cretaceous, although palaeobotanical and sedimentological evidences suggest that the climate was not uniform through the whole interval and that were two episodes in the Tithonian-Berriasian and Aptian-Albian periods of pronounced dry and/or arid environmental conditions. The composition and structure of the vegetation was not only affected by evolutionary changes but also by successive global geographical and climate changes. Hence, significant changes in the distribution of continental areas during the Mesozoic resulted in the latitudinal or sublatitudinal extensions of the climatic belts.     

Development and regional differentiation of the European vegetation during the Tertiary

The Tertiary vegetation of Europe evolved from paratropical to warm-temperate and temperate forms in response to a progressive, non-linear, climatic cooling. Its vegetational forms are composed mainly of two separate ecological units: the evergreen, laurophyll paleotropical geoflora and the deciduous, broad-leaved Arctotertiary geoflora. The development of the Tertiary climate and its interaction with the vegetation are convincingly indicated by the geoflora's migration; the changes in its composition; and the development of the Tertiary forest, swamp, and aquatic plant communities. The paleotropical geoflora is characterized in the upper Cretaceous to the upper Miocene by paratropical rain forest, subtropical rain and laurel forests, temperate laurel forests and edaphically-mediated formation of laurel-conifer forests. The Arctotertiary geoflora advanced into Europe in waves since the Paleocene and formed the basis for the Tertiary mixed mesophytic forests. These can be divided into warm-temperate rain forests, oak-hornbeamchestnut or mixed beech-oak-hornbeam forests, and edaphic formations such as bottomland and swamp forests. Beginning in the lower Cretaceous, the hydrophytic vegetation developed independently of the forest vegetation and formed very diverse herbaceous fresh water, swamp, salt water, and coastal formations. Considerable differences in composition allow to separate floral regions and provinces in Eurosiberia. Instead of three ill-defined floral regions in the Paleocene, there are four well-defined floral regions in the Pliocene. A Mediterranean region cannot be recognized, although Mediterranean (eumesogeic) floral elements appear in the Eocene/Oligocene and thereafter. The Mediterranean sclerophyll forests probably arose after the destruction of the laurophyll forests during the Pleistocene.     

Phylogeny of Acridocarpus-Brachylophon (Malpighiaceae): implications for tertiary tropical floras and Afroasian biogeography

Abstract.— A major tenet of African Tertiary biogeography posits that lowland rainforest dominated much of Africa in the late Cretaceous and was replaced by xeric vegetation as a response to continental uplift and consequent widespread aridification beginning in the late Paleogene. The aridification of Africa is thought to have been a major factor in the extinction of many African humid-tropical lineages, and in the present-day disparity of species diversity between Africa and other tropical regions. This primarily geologically based model can be tested with independent phylogenetic evidence from widespread African plant groups containing both humid- and xeric-adapted species. We estimated the phylogeny and lineage divergence times within one such angiosperm group, the acridocarpoid clade (Malpighiaceae), with combined ITS, ndhF , and trnL-F data from 15 species that encompass the range of morphological and geographic variation within the group. Dispersal-vicariance analysis and divergence-time estimates suggest that the basal acridocarpoid divergence occurred between African and Southeast Asian lineages approximately 50 million years ago (mya), perhaps after a southward ancestral retreat from high-latitude tropical forests in response to intermittent Eocene cooling. Dispersion of Acridocarpus from Africa to Madagascar is inferred between approximately 50 and 35 mya, when lowland humid tropical forest was nearly continuous between these landmasses. A single dispersal event within Acridocarpus is inferred from western Africa to eastern Africa between approximately 23 and 17 mya, coincident with the widespread replacement of humid forests by savannas in eastern Africa. Although the spread of xeric environments resulted in the extinction of many African plant groups, our data suggest that for others it provided an opportunity for further diversification.     

Classification and phytogeographical differentiation of oriental beech forests in Turkey and Bulgaria

Floristic differentiation of the oriental beech (Fagus orientalis Lipsky) forests in Turkey and Bulgaria was investigated and the role of geographical and topographical factors in this differentiation was assessed. After geographical and ecological stratification of the available 922 relevés, 288 remained. Classification, by applying cluster analysis, resulted in seven vegetation units defined by species composition which represent the geographical and ecological variation of Fagus orientalis forests. DCA ordination was applied to these units by passively projecting their chorological structure, as supplementary variables. For more detailed interpretation of vegetation types with similar geographic distribution patterns, PCA was applied by passively projecting the chorological elements, life-forms and topographical factors as supplementary variables. Seven vegetation units representing the geographical and ecological variety of Fagus orientalis forests were described. Four vegetation units represent the core area of Fagus orientalis distribution on the western and middle coast of the Black Sea region (Euxine region); the remaining three types represent the distribution in the eastern Black Sea region (Colchic region), the distribution in western and southern Anatolia under the influence of the Mediterranean climate and the distribution in the transitional zone from the Euxine region to the continental parts of Inner Anatolia, respectively. The four vegetation types in Euxine region reflect the decreasing effect of Black Sea towards Inner Anatolia, as well as altitudinal differences, except the forest type representing forests on calcareous sites. The other three vegetation units represent ravine, lowland to montane and altimontane forests in Euxine region. Fagus orientalis forests could be distinguished by their floristic composition, their chorological elements and life-forms spectra, which reflect a geographical and ecological gradients.     

太白山表土花粉和气孔器与植物类型的关系研究

通过陕西太白山13个样点表土花粉组合特征和气孔器及其与植物类型之间关系的分析,结果发现:针阔混交林花粉组合能很好地反映植物类型特征,落叶阔叶林和针叶林花粉组合能较好地与植物类型相对应,高山灌丛草甸花粉组合未能反映植物类型数量特征;主要花粉类型松属、铁杉属和桦属花粉具超代表性,胡桃属和榆属花粉具适宜代表性,落叶松属、冷杉属、杜鹃花科和槭属花粉具低代表性;DCA(Detrended Correspondence Analysis)分析表明,通过花粉数据能够较好区分不同植被类型,结合气孔器特征能够准确反映植被特征。     

History meets ecology: a geographical analysis of ecological restriction in the Neotropical humid montane forests avifaunas

Aim To analyse the ecological patterns of distribution of the avifaunas of the Neotropical humid montane forests, by assessing the degree of habitat restriction among species through the calculation of a numerical index, analysing their relationships with adjacent habitat, and exploring the relative contribution of both higher and lower habitats in shaping the avian assemblages. Location The Neotropical humid montane forests, from Mexico to north‐western Argentina. Methods The degree of species’ restriction to the habitat was calculated through a restriction index based on published endemicity indexes. The index scores range from 0 to 1: a score of 1 indicates a totally endemic species (i.e. fully restricted to the habitat); values tending to 0 indicate a widespread species. Results In Mesoamerica, completely restricted species represent a lower proportion of the total avifauna than in South American humid montane forests; whereas species shared to other habitats showed a higher proportion of the avifauna with affinities to higher altitude forests (e.g. pine and pine‐oak forests). South America, on the other hand, holds assemblages with a high proportion of completely restricted species; species shared to other habitats showed a high proportion of taxa with affinities to lowland forests. Main conclusions The ecological distribution of the HMF's avifauna could be partitioned in three main components: the ecologically restricted avifauna, the high altitude species and, the lower altitude species, which are tightly associated to the floristic composition along the gradient. The history of formation of the HMF flora and the ecological distribution of different avian taxa suggest a common history. Finally, the restriction index allowed a detailed evaluation of the composition of avian assemblages, their degree of restriction to the habitat and of the affinities regarding adjacent habitats, as well as an accurate distinction between species richness and restricted species richness, which should be a fundamental step towards the establishment of conservation priorities.     

Phytogeography of Italian deciduous oak woods based on numerical classification of plant distribution ranges

Abstract. 358 vascular plant species occurring in the deciduous oak woods of Italy were subdivided into 18 phytogeographical groups (chorotypes) through a numerical classification of their total ranges. The distribution patterns of the chorotypes are illustrated by chorograms, i.e. isopleth maps showing the joint distribution of species in Europe and surroundings regions. Except for Mediterranean species, which occur only occasionally in deciduous forests, the oak-wood flora of Italy consists of four main phytogeographical elements, which have a similar incidence (ca. 25 % each): (a) narrow-ranging, southern European species, (b) southeastern European species of submediterranean woodlands, (c) central-European species, (d) widely distributed European species. The oak woods of Italy differ from their central-European counterparts as to the relatively high floristic diversity, and in the presence of several narrow-ranging taxa, centred in different refugial areas for forest vegetation during the glacial period. There is a relation between the distribution patterns of the chorotypes and their relative frequencies in different communities; the latitudinal and longitudinal extensions of the ranges reflect the thermic and hygric conditions of the communities. The oak woods of northern and southern Italy are phytogeographically well-characterized; the former have a maximum of wide-ranging, central European species, the latter host several narrow-ranging taxa. The woodlands of central Italy have intermediate phytogeographical features, which is one of the main reasons for the difficulties in their syntaxonomical arrangement.     

Areas of high conservation value at risk by plant invaders in Georgia under climate change

Invasive alien plants (IAP) are a threat to biodiversity worldwide. Understanding and anticipating invasions allow for more efficient management. In this regard, predicting potential invasion risks by IAPs is essential to support conservation planning into areas of high conservation value (AHCV) such as sites exhibiting exceptional botanical richness, assemblage of rare, and threatened and/or endemic plant species. Here, we identified AHCV in Georgia, a country showing high plant richness, and assessed the susceptibility of these areas to colonization by IAPs under present and future climatic conditions. We used actual protected areas and areas of high plant endemism (identified using occurrences of 114 Georgian endemic plant species) as proxies for AHCV. Then, we assessed present and future potential distribution of 27 IAPs using species distribution models under four climate change scenarios and stacked single‐species potential distribution into a consensus map representing IAPs richness. We evaluated present and future invasion risks in AHCV using IAPs richness as a metric of susceptibility. We show that the actual protected areas cover only 9.4% of the areas of high plant endemism in Georgia. IAPs are presently located at lower elevations around the large urban centers and in western Georgia. We predict a shift of IAPs toward eastern Georgia and higher altitudes and an increased susceptibility of AHCV to IAPs under future climate change. Our study provides a good baseline for decision makers and stakeholders on where and how resources should be invested in the most efficient way to protect Georgia's high plant richness from IAPs.     

Quantitative phytogeography of the Italian Beech Forests

This paper presents a numerical analysis of the distribution patterns in Europe and adjoining regions of 242 vascular plant species occurring in the Italian beech forests. The classification of a matrix of species and of 531 Operational Geographic Units (OGUs) led to recognition of 15 different phytogeographic elements (chorotypes). The joint distribution of the species belonging to each chorotype is shown by a chorogram, which is a geographic map obtained by computer processing of the distributional data. Some of the chorograms show the main distributional centers for the beechwood flora in southern Europe: northwestern Balkan Peninsula and eastern Alps, southern Balkan Peninsula, Maritime Alps, northern Apennines, southern Apennines. Only 20% of the species are limited to narrow areas in southern Europe, and were not able to expand considerably their ranges in postglacial times. Migration from the refugia and colonization of vast areas in central and northern Europe led to intensive speciation phenomena, occurring chiefly through the formation of neopolyploid species. Several south European, narrow-ranging species, can be considered as palaeopolyploids with relict character. There is a good relationship between distribution and ecology of the chorotypes.Abbreviations OGU Operational Geographic Unit - OGS Operational Geographic Set - OCS Operational Character Set     

Trade-off between light availability and soil fertility determine refugial conditions for the relict light-demanding species in lowland forests

Identifying potential refugial habitats in the face of rapid environmental change is a challenge faced by scientists and nature conservation managers. Relict populations and refugial habitats are the model objects in those studies. Based on the example of Actaea europaea from Central Poland, we analyse the habitat factors influencing relict populations of continental, light-demanding species in lowland forests and examine which habitats of studied species corresponding most closely to ancient vegetation. Our results indicate that the current refugial habitats of Actaea europaea include not only communities which are very similar to ancient open forest but also forests with a closed canopy. Although the populations are influenced by nitrogen and light availability, the co-occurrence of these two factors in forest communities is limited by dense canopy formation by hornbeam and beech trees on fertile soils and in more humid conditions. Our findings indicate that the future survival of relict, light-demanding communities in lowland forests requires low-intensity disturbances to be performed in tree-stands, according to techniques, which imitate traditional forests management.     

Neotropical seasonally dry forests and Quaternary vegetation changes

Seasonally dry tropical forests have been largely ignored in discussions of vegetation changes during the Quaternary. We distinguish dry forests, which are essentially tree‐dominated ecosystems, from open savannas that have a xeromorphic fire‐tolerant, grass layer and grow on dystrophic, acid soils. Seasonally dry tropical forests grow on fertile soils, usually have a closed canopy, have woody floras dominated by the Leguminosae and Bignoniaceae and a sparse ground flora with few grasses. They occur in disjunct areas throughout the Neotropics. The Chaco forests of central South America experience regular annual frosts, and are considered a subtropical extension of temperate vegetation formations. At least 104 plant species from a wide range of families are each found in two or more of the isolated areas of seasonally dry tropical forest scattered across the Neotropics, and these repeated patterns of distribution suggest a more widespread expanse of this vegetation, presumably in drier and cooler periods of the Pleistocene. We propose a new vegetation model for some areas of the Ice‐Age Amazon: a type of seasonally dry tropical forest, with rain forest and montane taxa largely confined to gallery forest. This model is consistent with the distributions of contemporary seasonally dry tropical forest species in Amazonia and existing palynological data. The hypothesis of vicariance of a wider historical area of seasonally dry tropical forests could be tested using a cladistic biogeographic approach focusing on plant genera that have species showing high levels of endemicity in the different areas of these forests.     

Early Cretaceous angiosperm invasion of Western Europe and major environmental changes

BACKGROUND AND AIMS: At the beginning of the Late Cretaceous, angiosperms already inhabited all the environments and overtopped previously gymnosperm-dominated floras, especially in disturbed freshwater-related environments. The aim of this paper is to define what fossil plant ecology occurred during the early Cretaceous in order to follow the early spread of angiosperm taxa. METHODS: Floristic lists and localities from the Barremian to the Albian of Europe are analysed with the Wagner's Parsimony Method. KEY RESULTS: The Wagner's Parsimony Method indicates that (a) during the Barremian, matoniaceous ferns formed a savannah-like vegetation, while angiosperms composed freshwater aquatic vegetation; (b) during the Late Aptian humid phase, conifers increased, while matoniaceous ferns decreased, reflecting the closure of the vegetation; and (c) from the Albian, warmer and drier conditions induced the recovery of the matoniaceous ferns, while core angiosperms first developed in floodplains. CONCLUSIONS: During the late Early Cretaceous (Barremian-Albian), angiosperms showed a stepwise widening of their ecological range, being recorded first during the Barremian as aquatic plant mega-remains and at the Cenomanian onwards occurred in all the environments.     

Effects of altitude and climate in determining elevational plant species richness patterns: A case study from Los Tuxtlas,Mexico

Altitudinal changes of composition and richness of montane plant assemblages are complex, depending on the taxonomic group and gradient conditions, with different factors involved that are directly altitude-dependent (e.g., temperatures, air pressure) and altitude-independent (e.g., precipitation, cloud cover, area). In order to assess the relative impacts of temperature, precipitation, air humidity, and area of altitudinal belts on plant diversity, we analyzed diversity patterns of five species-rich groups, mostly herbaceous plants, in 74 forest plots along three climatically contrasting elevational transects from humid tropical lowland vegetation up to cloud forests at Los Tuxtlas, Mexico. We recorded 278 plant species, with ferns being the most species-rich group followed by orchids, bromeliads, aroids, and piperoids. The most striking results were the contrasting patterns and model results for terrestrial and epiphytic taxa. Whereas the richness of all terrestrial species taken together did not change significantly with elevation, vascular epiphytes showed increasing species numbers with altitude. However, a number of individual terrestrial taxa showed also significant elevation-related changes: aroids showed a marked decline with hight, orchids and piperoids increased, and ferns displayed a hump-shaped pattern with highest richness in mid-altitudes. Among the epiphytes, aroids declined while most other groups increased with altitude. This distinction is relevant for projections of responses of plant communities to climate change, which will lead to increased temperatures and to changing precipitation and cloud condensation regimes and thus will likely affect terrestrial and epiphytic species in different ways.     

Climate and vegetational regime shifts in the late Paleozoic ice age earth

The late Paleozoic earth experienced alternation between glacial and non-glacial climates at multiple temporal scales, accompanied by atmospheric CO₂ fluctuations and global warming intervals, often attended by significant vegetational changes in equatorial latitudes of Pangaea. We assess the nature of climate–vegetation interaction during two time intervals: middle–late Pennsylvanian transition and Pennsylvanian–Permian transition, each marked by tropical warming and drying. In case study 1, there is a catastrophic intra-biomic reorganization of dominance and diversity in wetland, evergreen vegetation growing under humid climates. This represents a threshold-type change, possibly a regime shift to an alternative stable state. Case study 2 is an inter-biome dominance change in western and central Pangaea from humid wetland and seasonally dry to semi-arid vegetation. Shifts between these vegetation types had been occurring in Euramerican portions of the equatorial region throughout the late middle and late Pennsylvanian, the drier vegetation reaching persistent dominance by Early Permian. The oscillatory transition between humid and seasonally dry vegetation appears to demonstrate a threshold-like behavior but probably not repeated transitions between alternative stable states. Rather, changes in dominance in lowland equatorial regions were driven by long-term, repetitive climatic oscillations, occurring with increasing intensity, within overall shift to seasonal dryness through time. In neither case study are there clear biotic or abiotic warning signs of looming changes in vegetational composition or geographic distribution, nor is it clear that there are specific, absolute values or rates of environmental change in temperature, rainfall distribution and amount, or atmospheric composition, approach to which might indicate proximity to a terrestrial biotic-change threshold.     

Vegetation patterns and their dependency on site conditions in the pre-industrial landscape of north-eastern Japan

We reconstructed the vegetational landscape of the pre-industrial era (the beginning of the twentieth century) in north-eastern Japan, and estimated the distribution patterns of traditional land-uses, as suggested from the vegetation. We found significant correspondence between the spatial patterns of vegetational landscape and site attributes, and hypothesized the underlying mechanisms. The study area was classified into three vegetation types: grasslands, secondary forests and old-growth forests. It was determined that the grasslands were formed and maintained by burning; secondary forests were derived from either charcoal woods or forests recovered on abandoned grasslands; and old-growth forests had suffered the least anthropogenic disturbance. Each past vegetation type showed significant dependency on site attributes such as altitude, slope angle, slope aspect, hydrological topography and distance from the nearest human habitation. The relative importance of these site attributes varied depending on the vegetation type. Grasslands and old-growth forests, which were the most and the least disturbed sites in the study area, respectively, showed clear contrasts in their dependencies especially on the slope aspect and on elevation. These site attributes were thought to have had influences on each vegetation type by determining the inflammability of the site. Satellite photographs indicated that north-facing valleys had been relatively wet throughout the fire-prone spring season. Hence, these areas would have been free from frequent fire, and more likely to preserve old-growth forests. Ground wetness in spring was thought to be the underlying factor determining the contrasts in past vegetation and land-use patterns in the area.An erratum to this article can be found at     

European post-glacial forests: compositional changes in response to climatic change

Abstract. Multivariate analysis of an extensive palyno-logical database for Europe has enabled reconstruction of broad-scale vegetation history. Whereas many major features of present vegetation patterns were established early in the Holocene, floristic composition of the forests has changed continuously up to the present day. For example, although ‘mixed deciduous forests’ had reached approximately their present extent in northwest Europe by 8000 B.P., Tilia peaked in abundance in these forests during the middle post glacial, whereas Pinus was most abundant in these forests during the early post-glacial and Fagus increased in abundance only in recent millennia. Pollen-climate response surfaces for major pollen taxa show how their distribution and abundance patterns relate to contemporary climate. Past forest-compositional changes were responses to climatic changes, the nature of which can be inferred from pollen-climate response surfaces. Post-glacial climate changes have been different in magnitude and direction in different regions of Europe. For example, in recent millennia the vegetation changes indicate decreasing summer temperatures in northern Europe but increasing summer temperatures in the Mediterranean region. The way in which vegetation responded to past climatic changes gives insight into the likely response of vegetation to future climate changes induced by the ‘greenhouse effect’.     

Zelkova carpinifolia (Pallas) K. Koch in Holocene sediments of Georgia—an indicator of climatic optima

Zelkova pollen has been found in Oligocene- to Pleistocene-aged deposits from many parts of Europe and northern Africa, but became extinct in mainland Europe prior to the last glacial maximum. This paper presents some observations on the ecology, pollen productivity and Holocene history of Zelkova carpinifolia to further understanding of Quaternary climatic trends. Georgia is one of the last refuges of this Tertiary relict tree.Based on palynological data from 20 Holocene sediment profiles in Georgia, we have established that Zelkova pollen is almost always accompanied by elevated proportions of thermophilous taxa (Castanea sativa, Quercus hartwissiana, Quercus iberica, and Pterocarya pterocarpa) in pollen spectra. These spectra are associated with phases of climatic amelioration and humidification. Zelkova carpinifolia is characterised by low pollen productivity and is underrepresented in pollen spectra by four-to-five orders of magnitude. Because of this, even single grains of Zelkova pollen may play a significant role in pollen-based climatic reconstructions.Six major climatic optima occurred in Georgia through the course of the Holocene, the longest and warmest of which occurred in the mid-Holocene and reached its maximum between 6000 and 5500 Cal. yr BP. During that period, Zelkova and Castanea forests were widespread. In Western Georgia, the upper tree line was elevated by as much as 300 m above its present-day level. In semiarid Eastern Georgia, the tree line may have been 500–600 m higher. Other climatic optima are seen in late-Holocene pollen spectra dating to 3800–2500 Cal. yr BP and 1350–800 Cal. yr BP.