Ecosystems are defined by the community of living organisms and how they interact together and with theirenvironment. Insects and plants are key taxa in terrestrial ecosystems and their network determines the trophicstructure of the environment. However, what drives the interactions between plants and insects in modern andfossil ecosystems is not well understood. In this study, we analyzed insect damage richness and frequency in 5000 fossil leaves deposited during the early Miocene at 20–17 Ma along a latitudinal gradient from Europe (twolocalities in Czech Republic) to Turkey (one locality) in a temperate climate setting. Damage frequency wasmainly linked with abiotic factors (temperature, precipitation seasonality) whereas damage richness was mainlylinked with biotic factors (plant richness, biome). Univariate analysis of insect damage types consistently suggested closer trophic similarity between the Mediterranean and either the one or the other Central European plant assemblage. In contrast, multivariate analysis of all insect damage types indicated closer similarity between the two Central European sites highlighting the importance of biogeographic legacy and geographic closeness to the plant-insect interaction patterns. Our results underscore the high complexity of the herbivory network andcall for careful interpretations of plant-insect interaction patterns in palaeoecological studies. Finally, comparing the trophic similarity between different localities using total evidence plots as done in this work might be apromising complementary method in comparative studies of plant-insect interactions.

The uppermost Eocene Florissant Formation, Rocky Mountains, Colorado, has yielded numerous insect, vertebrate, and plant fossils. Three previous comprehensive palynological studies investigated sections of lacustrine deposits of the Florissant Formation and documented the response of plant communities to volcanic eruptive phases but overall found little change in plant composition throughout the investigated sections. These studies reported up to 150 pollen and spore phenotypes. In the present paper we used a taxonomic approach to the investigation of dispersed pollen and spores of the Florissant Formation. Sediment samples from the shale units containing macrofossils were investigated using light microscopy (LM) and scanning electron microscopy (SEM). The general picture of the palynoflora is in agreement with previous studies. However, the combined LM and SEM investigation provides important complementary information to previous LM studies. While a fairly large amount of previous pollen determinations could be confirmed, the purported taxonomic affinities of several pollen phenotypes need to be revised. For example, pollen referred to as Podocarpus or Podocarpidites sp. belongs to the Pinaceae Cathaya, Malus/Pyrus actually belongs to Dryadoideae, pollen of the form genus Boehlensipollis referred to as Proteaceae/Sapindaceae/Elaeagnaceae or Cardiospermum belongs to Sapindaceae but not to Cardiospermum, and pollen of Persicarioipollis sp. B with previously assumed affinities to Polygonaceae actually belongs to Thymelaeaceae. Pandaniidites and one type of Malvacipollis cannot be linked with Pandanaceae and Malvaceae. A few taxa are new records for Florissant (Ebenaceae: Diospyros; Mernispermaceae; Trochodendraceae: Tetracentron). In general, SEM investigations complement the LM palynological studies and improve the identification of dispersed pollen and spores and enable integration of data from dispersed fossil pollen into a wide range of comparative morphological, taxonomic, evolutionary, biogeographic, and phylogenetic studies.

The piggyback basin of Bezhan, southeastern Albania, was formed during the late Neogene and contains Pliocene/Pleistocene deposits. These continental deposits consist of marls, siltstones and clays separated by a thin series of lignite-seams alternating with clays (Bezhan formation). We investigated leaf fossils and dispersed pollen from marls of the upper portion of this formation. Fifty-two plant taxa comprising algae, gymnosperms, and angiosperms were recovered. Of these, at least 19% belong to extant species and less than 16% belong to taxa today extinct in western Eurasia. Tsuga is represented by three pollen taxa with affinities to modern Chinese, Japanese, and North American species. Herbaceous taxa indicative of steppe (Artemisia, Amaranthaceae) occur in low quantities (≤1%) suggesting an interglacial setting. Four vegetation units are recognised: Wet riparian and aquatic vegetation, mesic oak forest, dry sub-Mediterranean woodland, and montane conifer forest. A comparison of the Bezhan flora with well-dated Pliocene and Pleistocene floras of Italy suggests a Calabrian (late early Pleistocene) age for the upper unit. This estimate is based on the abundance of extant taxa, the absence of subtropical taxa, and threshold values of particular taxa (Tsuga, Carya). The findings are in agreement with age estimates for extant tree species from molecular studies.

The former family Taxodiaceae is currently treated as nine genera in five subfamilies of the family Cupressaceae. Pollen ofthe ‘taxodiaceous’ Cupressaceae typically has a papilla in the leptoma area and is common in Cenozoic strata because someof its genera were key elements in lignite forming swamp forests. Dispersed fossil pollen of this group are often assigned toparticular genera and, based on the modern ecologies of these taxa, to particular palaeoenvironments. In this study, weinvestigated pollen of all nine genera of the former Taxodiaceae using light and scanning electron microscopy to evaluatewhether pollen morphology can be used to discriminate modern genera of this paraphyletic group. We found few genus- orsubfamily-diagnostic characters among members of taxodiaceous Cupressaceae. Features such as orbiculae and pollen andleptoma size cannot be used to discriminate subfamilies. However, three basal subfamilies share short papillae, whereas intwo more derived clades (Sequoioideae and Taxodioideae) papillae are markedly longer. In the generally non-papillate coreCupressaceae, the leptoma (aperture) area may or may not possess a distinct circular thinning as also found in the basalgrade of taxodiaceous Cupressaceae. Our results show that it is difficult if not impossible to distinguish genera of theecologically distinct Taxodioideae and Sequoioideae based on pollen morphology. In view of a much wider ecologicalamplitude of many taxodiaceous Cupressaceae during large parts of the Cenozoic, we conclude that it is not recommendableto infer particular palaeoenvironments on the basis of dispersed taxodiaceous pollen grains alone.

The excavated main lignite seams and overlying lacustrine sediments of the opencast mines Eskihisar, Salihpaşalar, and Tınaz, Muğla Province, south-western Turkey were investigated using a high taxonomic resolution palynological approach.

The Eskihisar section comprises 47m and 56 samples of which 30 were usable for palynological analysis. The Tınaz section comprises 75 m and 29 samples of which 15 were usable for palynological analysis. Finally, the Salihpaşalar section comprises 25 m and 26 samples of which 16 were usable for palynological analysis. The age of the palynological sections is middle to late Miocene based on radiometric dating and vertebrate fossils.

In order to investigate dispersed pollen and spores and their botanical affinities a combined light microscopy and scanning electron microscopy approach was used. The rich palynoflora comprises seven types of algal cysts (Botryococcus, Zygnemataceae), seventeen spore types (Lycopsida, Marsileaceae, Osmundaceae, Pteridaceae, Polypodiaceae), 14 types of gymnosperm pollen (Ephedraceae, Cupressaceae, Pinaceae), five types of monocotyledone pollen (Poaceae, Typhaceae) and ca 90 dicotyledone pollen types (Altingiaceae, Amaranthaceae, Anacardiaceae, Apiaceae, Aquifoliaceae, Asteraceae, Betulaceae, Campanulaceae, Cannabaceae, Caprifoliaceae, Caryophyllaceae, Ericaceae, Eucommiaceae, Euphorbiaceae, Fabaceae, Fagaceae, Geraniaceae, Juglandaceae, Lamiaceae, Linaceae, Lythraceae, Malvaceae, Myricaceae, Oleaceae, Onagraceae, Plumbaginaceae, Polygonaceae, Ranunculaceae, Rosaceae, Salicaceae, Sapindaceae, Sapotaceae, Ulmaceae).

The objectives of this investigation were (1) to evaluate whether the three palynological sections were deposited at the same time, and (2) to show regional vegetation differences within a single sedimentary basin.

We found three general pollen zones corresponding to different sedimentary settings and palaeoenvironments. The first pollen zone was linked to lignite formation (swamp forest, fern spores, Alnus, Decodon). The second pollen zone reflects lacustrine conditions (Typhaceae) and surrounding hinterland vegetation dominated by Fagaceae. The third pollen zone is dominated by herbaceous taxa, whereas woody taxa are less diverse and less abundant.

In general, the three palynological sections are congruent in reflecting distinct pollen zones. However main vegetation types may be represented by different dominating taxa (e. g. Alnus dominace in Eskihisar and Tınaz localities while absent in Salihpaşalar) and rare taxa may differ between localities.

Our results demonstrate that in order to achieve a comprehensive understanding of environmental and vegetation conditions in a sedimentary basin, a single palynological section (locality) may not capture the entirety of environmental conditions and changes.

Anatolia was a crossroads for mammal migration during the Miocene due to intermittent land connections between Africa and Anatolia and persisting warm conditions. Here, we investigated a palynological section from middle Miocene sediments of Eskihisar (southwestern Anatolia) in order to establish biogeographic links of the palynoflora and to infer the palaeoenvironment. Four algal palynomorphs, nine spore taxa, eight gymnosperms, three monocots, and 67 dicot pollen types were encountered and investigated using the “single grain method” that combines light microscopy and scanning electron microscopy. Two pollen zones reflect different phases of basin development. Zonal vegetation remained fairly stable across the section and reflects heterogeneous environments including broad-leaved deciduous forest, subtropical forest, and sclerophyllous and semi-evergreen oak forest. Conifers were accessory elements in the broad-leaved deciduous forest communities and replaced these at higher elevations. Some herbaceous taxa (Plumbaginaceae) indicate scattered occurrences of sandy and/or rocky soils. Biogeographic affinities are general Northern Hemispheric, North American, and East Asian as also suggested by the macro fossil record. Only two taxa provide potential biogeographic links with the African flora. This suggests that biome shifts of plant taxa between African subtropical /tropical biomes and Anatolian (western Eurasian) temperate forests and shrublands may have been rare in the middle Miocene.

In an ongoing study, focussing on the plant fossils and palynofloras of the lignite strip mines of the Yatağan basin(Muğla province), a number of pollen taxa, previously not reported from middle Miocene terrestrial sediments of Anatolia were encountered.

The middle Miocene climate transition (MMCT) was a phase of global cooling possibly linked to decreasing levels of atmospheric CO2. The MMCT coincided with the European  Mammal Faunal Zone MN6. From this time, important biogeographic links between Anatolia  and eastern Africa include the hominid Kenyapithecus. Vertebrate fossils suggested mixed  open and forested landscapes under (sub)tropical seasonal climates for Anatolia. Here, we  infer the palaeoclimate during the MMCT and the succeeding cooling phase for a middle Miocene (14.8–13.2 Ma) of an intramontane basin in southwestern Anatolia using three2palaeobotanical proxies: (i) Köppen signatures based on the nearest-living-relative principle. (ii) Leaf physiognomy analysed with the Climate Leaf Analysis Multivariate Program (CLAMP). (iii) Genus-level biogeographic affinities of fossil floras with modern regions. The three proxies reject tropical climates for the MMCT of southwestern Anatolia and instead infer warm temperate C climates. Köppen signatures reject summer-dry Cs climates but cannot discriminate between fully humid Cf and winter-dry Cw; CLAMP reconstructs Cf climate based on the low X3.wet/X3.dry ratio. Additionally, we assess whether the palaeobotanical record does resolve transitions from the warm Miocene Climatic Optimum (MCO, 16.8–14.7 Ma) into the MMCT (14.7–13.9 Ma), and a more pronounced cooling at 13.9–13.8 Ma, as reconstructed from benthic stable isotope data. For southwestern Anatolia, we find that arboreal taxa predominate in MCO floras (MN5), whereas in MMCT floras (MN6) abundances of arboreal and non-arboreal elements strongly fluctuate indicating higher structural complexity of the vegetation. Our data show a distinct pollen zone between MN6 and MN7+8 dominated by herbaceous taxa. The boundary MN6 and MN7+8, roughly corresponding to a first abrupt cooling at 13.9–13.8 Ma, possibly might be associated with this herb-rich pollen zone.

The subject of this study is the palynology (biostratigraphic and taxonomic) and the plant remains of the lignite strip mines of Eskihisar, Salihpasalar, and Tinaz (Muğla province, western Turkey). In the Yatağan basin two Miocene to Pliocene formations are present, the Eskihisar Formation (early to middle Miocene) and the Yatağan Formation (late Miocene to early Pliocene). Both formations represent river and lake deposits consisting mainly of conglomerate, sandstone, claystone, limestone, tuffite, and intercalated lignite; the thickest, actively mined lignite seams occur in the Sekköy member of the Eskihisar Formation.

Previous palynological studies of the palynoflora of the Yatağan basin mainly focussed on its biostratigraphic and palaeoclimatic significance, using conventional morphological nomenclature and light microscopy (LM).

In this study the „single grain method“ is applied. Using this method, the same individual pollen grains are investigated by using both LM and scanning electron microscopy (SEM). The resulting high-resolution pictographs enable a much higher taxonomic resolution.

The studied palynoflora is very rich and taxonomically diverse. Cryptogams are represented by more than ten spore morphotypes of at least three families (Osmundaceae, Pteridaceae, Polypodiaceae). Gymnosperm pollen is dominated by Cupressaceae, Gnetales (Ephedra), and Pinaceae (Cathaya, Keteleeria, Pinus). Angiosperm pollen can be assigned to 57 different genera belonging to Poaceae, Typhaceae, Altingiaceae, Amaranthaceae (Chenopodieae), Anacardiaceae, Apiaceae (three types), Asteraceae (Asteroideae, Cichoriodeae), Betulaceae (Alnus, Betula, Carpinus, Ostrya) Buxaceae, Campanulaceae, Caprifoliaceae (Lonicera), Caryophyllaceae, Dipsacaceae, Eucommiaceae, Euphorbiaceae, Fabaceae, Fagaceae (Fagus, Quercus, Trigonobalanopsis) Geraniaceae, Juglandaceae, Linnaceae (Linnum), Malvaceae, Myricaceae, Nymphaeaceae, Oleaceae (four different types), Plumbaginaceae (Armeria,), Polygonaceae (Rumex), Rosaceae, Sapindaceae (Acer), Ulmaceae (Cedrelospermum, Ulmus, Zelkova), and Zingiberales (Spirematospermum).

In addition, more than two thousand plant macrofossils were collected in the course of repeated field trips, including remains of Pinaceae, Berberidiaceae (Mahonia), Betulaceae (Alnus, Carpinus), Buxaceae (Buxus), Fagaceae (Fagus, Quercus), Lauraceae, Malvaceae (Tilia), Myricaceae (Myrica), Rosaceae, Salicaceae (Populus, Salix), Sapindaceae (Acer), Smilacaceae (Smilax), Typhaceae (Typha), Ulmaceae (Zelkova).

A combined analysis integrating these rich and diverse plant macro- and microfossil records will lead to a better understanding and refined reconstruction of the vegetation in the Yatağan basin during the middle to late Miocene.

The late Miocene is marked by pronounced environmentalchanges and the appearance of strong temperature andprecipitation seasonality. Although environmental heterogeneityis to be expected during this time, it is challenging to reconstructpalaeoenvironments using plant fossils. We investigated leavesand dispersed spores/pollen from 6.4 to 6 Ma strata inthe intermontane Florina–Ptolemais–Servia Basin (FPS) ofnorthwestern Greece. To assess how well plant fossils reflectthe actual vegetation of the FPS, we assigned fossil taxa tobiomes providing a measure for environmental heterogeneity.Additionally, the palynological assemblage was compared withpollen spectra from modern lake sediments to assess biases inspore/pollen representation in the pollen record. We found aclose match of the Vegora assemblage with modern Fagus–Abiesforests of Turkey. Using taxonomic affinities of leaf fossils, wefurther established close similarities of the Vegora assemblage with modern laurophyllous oak forests of Afghanistan. Finally, using information from sedimentaryenvironment and taphonomy, we distinguished local and distantly growing vegetation types.We thensubjected the plant assemblage of Vegora to different methods of climate reconstruction and discussedtheir potentials and limitations. Leaf and spore/pollen records allow accurate reconstructions ofpalaeoenvironments in the FPS, whereas extra-regional vegetation from coastal lowlands isprobably not captured.

Middle Miocene deposits exposed at the Tınaz lignite mine, Yatağan Basin, Muğla, southwestern Turkey, were palynologically investigated. The Tınaz lignite mine section belongs to the Eskihisar Formation. The lignite seam at the base of the section represents the uppermost part of the Turgut Member. Above, c. 65 m of clayey siltstone, limestone, and marls represent the Sekköy Member. Nine spores, zygospores and cysts of fungi and algae, seven moss and fern spores, 12 gymnosperm pollen types, and more than 80 angiosperm pollen taxa were recovered from the Tınaz lignite mine section. Three pollen zones were recognized, of which pollen zone 1 corresponds to the formation of the main lignite seam and reflects the change from a fluviatile to a lacustrine depositional setting. Pollen zones 2 and 3 and a transitional zone 2-3 reflect different stages of lake development and a shift in local vegetation from forested (pollen zones 1 and 2) to more open (transitional zone 2-3, zone 3). Interpreting changes in regional vegetation from pollen zones 1 to 3 is not straightforward as changes in the pollen spectra may be affected by changing contributions of airborne and water transported pollen and spores to the observed palynoassemblages. Age inference for the Tınaz lignite mine section has been complicated by the absence of datable ash layers, associated mammal faunas, or marine sediments. However, pollen zone 3 shares key features with the pollen spectrum recovered from the nearby mammal site Yenieskihisar (upper part of Sekköy Member) for which an age of 12.5-11.2 Ma has been suggested, and to the youngest pollen zone recovered from the mammal locality Çatakbağyaka, 10 km south of Tınaz, that probably represents mammal zone MN7/8 instead of MN5 or MN6 as previously suggested. In contrast, pollen zones 1 and 2 are fairly similar to the basal parts of the Çatakbağyaka pollen flora (uppermost parts of Turgut Member, basalmost parts of Sekköy Member). Furthermore, new mammal data from the Yatağan basin suggest that the layers below pollen zone 1 are MN4/5, and that carnivores cooccuring with pollen zone 1 in the main lignite seam of Eskihisar probably belong to MN6. Hence, a Langhian to Serravallian age can be inferred for pollen zones 1 and 2 of the Tınaz lignite mine section, and a late Serravallian age for pollen zone 3. Palaeobiogeographic relationships of the palynofloras are generally northern hemispheric, with many north temperate tree taxa showing modern disjunctions East Asia- NorthAmerica (Tsuga, Carya), East Asia- western Eurasia (Zelkova), East Asia- North America- western Eurasia (Liquidambar), or restricted to East Asia (Cathaya, Eucommia) or North America (Decodon). A few taxa belong to extinct lineages that have complex biogeographic patterns (Engelhardioideae, Cedrelospermum). The presence of Picrasma (Simaroubaceae) in the lower lignite layers of pollen zone 1 is remarkable, as the botanical affinities with the enigmatic flower Chaneya present in early to middle Miocene deposits of Turkey and Central Europe have recently been shown to be with Picrasma. 

Standard models of plant speciation assume strictly dichotomous genealogies in which a species, theancestor, is replaced by two offspring species. The reality in wind-pollinated trees with long evolutionaryhistories is more complex: species evolve from other species through isolation when genetic drift exceeds gene flow; lineage mixing can give rise to new species (hybrid taxa such as nothospecies and allopolyploids). The multi-copy, potentially multi-locus 5S rDNA is one of few gene regions conserving signal from dichotomous and reticulate evolutionary processes down to the level of intra-genomic recombination. Therefore, it can provide unique insights into the dynamic speciation processes of lineages that diversified tens of millions of years ago. Here, we provide the first high-throughput sequencing (HTS) of the 5S intergenic spacers (5S-IGS) for a lineage of wind-pollinated subtropical to temperate trees, the Fagus crenata – F.sylvatica s.l. lineage, and its distant relative F. japonica. The observed 4963 unique 5S-IGS variants reflect acomplex history of hybrid origins, lineage sorting, mixing via secondary gene flow, and intra-genomic competition between two or more paralogous-homoeologous 5S rDNA lineages. We show that modern species are genetic mosaics and represent a striking case of ongoing reticulate evolution during the past 55 million years.

Species distribution models can help predicting range shifts under climate change. The aimof this study is to investigate the late Quaternary distribution of Oriental beech (Fagus orientalis)and to project future distribution ranges under different climate change scenarios usinga combined palaeobotanical, phylogeographic, and modelling approach. Five species distributionmodelling algorithms under the R-package ‘biomod2‘were applied to occurrence dataof Fagus orientalis to predict distributions under present, past (Last Glacial Maximum, 21ka, Mid-Holocene, 6 ka), and future climatic conditions with different scenarios obtainedfrom MIROC-ESM and CCSM4 global climate models. Distribution models were comparedto palaeobotanical and phylogeographic evidence. Pollen data indicate northern Turkey andthe western Caucasus as refugia for Oriental beech during the Last Glacial Maximum.Although pollen records are missing, molecular data point to Last Glacial Maximum refugiain northern Iran. For the mid-Holocene, pollen data support the presence of beech in thestudy region. Species distribution models predicted present and Last Glacial Maximum distributionof Fagus orientalis moderately well yet underestimated mid-Holocene ranges.Future projections under various climate scenarios indicate northern Iran and the Caucasusregion as major refugia for Oriental beech. Combining palaeobotanical, phylogeographicand modelling approaches is useful when making projections about distributions of plants.Palaeobotanical and molecular evidence reject some of the model projections. Nevertheless,the projected range reduction in the Caucasus region and northern Iran highlights theirimportance as long-term refugia, possibly related to higher humidity, stronger environmentaland climatic heterogeneity and strong vertical zonation of the forest vegetation.

Diospyros is a large genus of woody flowering plants with a predominantly subtropical and tropical modern distribution. Fossils attributed to Diospyros are mainly leaf impressions from Cretaceous and Cenozoic strata across the Northern Hemisphere. However, it is difficult to assign such fossils to Diospyros because genus-diagnostic leaf characteristics remain to be identified. Unequivocal fossil records of Diospyros are few, including staminate flowers with in situ pollen from the late Eocene of South Australia and dispersed pollen from Cenozoic strata in the Northern Hemisphere. Here, we investigated dispersed pollen and calyx remains from Miocene deposits of Denmark using a combined scanning electron/light microscopy approach. Tricolporate, relatively large pollen with lalongate pori and longbow-shaped colpi and a distinctive micro/nanorugulate exine ornamentation together with persistent 4-lobed flower calyces allow unambiguous identification of the genus. Based on the large size of the calyx, we describe a new fossil-species of Diospyros. Further, a review of the fossil pollen record of Diospyros shows that, in addition to the Australian record, the genus was present in South China, western North America and Europe during the Eocene and in East and South Africa and Central Asia during the Oligo-Miocene. Although still scanty, the pollen record can contribute vital information for time calibrated molecular phylogenies to resolve conflicting biogeographic inferences. A thorough description of the historical biogeography of Diospyros is still in its infancy. While we initiate such a study here, development of a comprehensive picture will require further studies of dispersed pollen grains with high taxonomic resolution.

Premise: The Fagaceae comprise around 1000 tree species in the Northern Hemisphere.Despite an extensive fossil pollen record, reconstructing biogeographic patternsis hampered because it is difficult to achieve good taxonomic resolution withlight microscopy alone. We investigate dispersed pollen of Fagaceae from the Miocene Søby flora, Denmark. We explore the latitudinal gradient in Fagaceae distribution during the Miocene Climatic Optimum (MCO) in Europe and the Northern Hemisphere to compare it with the Eocene Warmhouse and the present.

Methods: We investigated dispersed pollen using light and scanning electron microscopy. We assessed biogeographic patterns in Fagaceae during two warm periods in Earth history (MCO, Eocene) and the present.

Results: Eight species of Fagaceae were recognized in the Søby flora. Of these, Fagus had a continuous Mediterranean to subarctic distribution during MCO; Quercus sect. Cerris and castaneoids had northern limits in Denmark, and evergreen Quercus sect. Ilex in Central Europe. In a northern hemispheric context, Fagus and sections of Quercus had more northerly distribution limits during Eocene and MCO with maximum northward extensions during Eocene (Fagus, castaneoids) or Oligo‐Miocene (Quercus sects. Cerris and Ilex). The known distribution of the extinct Tricolporopollenites theacoides during MCO included Central Europe and East China, while this taxon thrived in South China during Eocene.

Conclusions: More northerly distributions during MCO and Eocene probably were determined by temperature. In contrast, fossil occurrences in areas that are arid or semi‐humid today were determined by maritime conditions in these areas (western North America, Central Asia) during the Cenozoic.

Previous paleobotanical work concluded that Paleogene elements of the sclerophyllous subhumid vegetation of western Eurasia and western North America were endemic to these disjunct regions, suggesting that the southern areas of the Holarctic flora were isolated at that time. Consequently, molecular studies invoked either parallel adaptation to dry climates from related ancestors, or long-distance dispersal in explaining disjunctions between the two regions, dismissing the contemporaneous migration of dry-adapted lineages via land bridges as unlikely.

We report Vauquelinia (Rosaceae), currently endemic to western North America, in Cenozoic strata of western Eurasia. Revision of North American fossils previously assigned to Vauquelinia confirmed a single fossil-species of Vauquelinia and one of its close relative Kageneckia.

We established taxonomic relationships of fossil-taxa using diagnostic character combinations shared with modern species and constructed a time-calibrated phylogeny.

The fossil record suggests that Vauquelinia, currently endemic to arid and subdesert environments, originated under seasonally arid climates in the Eocene of western North America and subsequently crossed the Paleogene North Atlantic land bridge (NALB) to Europe. This pattern is replicated by other sclerophyllous, dry-adapted and warmth-loving plants, suggesting that several of these taxa potentially crossed the North Atlantic via the NALB during Eocene times.

Late Oligocene leaf assemblages from four sites in Southwestern Siberia (Kurgan, Tyumen, Omsk oblasts) are described. Twenty-three leaf taxa and 3 reproductive structures represent local vegetation of a lake (Salvinia,Typha, Phragmites, Nelumbo, Hemitrapa, Liquidambar, Pterocarya, Alnus, Populus, Salix, Nyssa). Additionally, 57 spore and pollen taxa were recorded from one site (Shish River). Gymnosperms dominate the assemblage with ~30% Pinaceae and ~25% taxodiaceous (papillate) Cupressaceae pollen. Ferns and peat mosses (Sphagnaceae) comprised ~6%. Angiosperms were dominated by Fagaceae, Betulaceae, Juglandaceae and Ulmus and comprised a few exotic elements (Liquidambar, Eucommia, Nyssa, Symplocos); scarce herbaceous plants reflect lakeshore vegetation. The flora of the Turgay type comprised old elements (Nelumbo protospeciosa, Liquidambar europaea, taxodiaceous/papillate Cupressaceae, Quercus sect. Protobalanus) and taxa present in Siberia/Kazakhstan during the Paleogene with later arrivals in Europe (Ulmus pyramidalis, Quercus pseudocastanea, Alnus julianiformis, Byttneriophyllum tiliifolium). A few taxa were endemic in the late Oligocene of western Siberia (Trapa praeconocarpa, Platycrater iljinskajae sp. nov.). Combined macrofossil and palynological evidence places the Shish River site flora into the late Oligocene Zhuravka (Turtas) Formation. Floras of similar composition from western Eurasia are commonly middle Miocene or youngerin age highlighting the dynamic spatiotemporal evolution of temperate Eurasian floras during the Cenozoic.

Background and Aims: Cork oaks (Quercus section Cerris) comprise 15 extant species in Eurasia. Despite being a small clade, they display a range of leaf morphologies comparable to the largest sections (>100 spp.) in Quercus. Their fossil record extends back to the Eocene. Here, we explore how cork oaks achieved their modern ranges and how legacy effects might explain niche evolution in modern species of section Cerris and its sister section Ilex, the holly oaks.

Methods: We inferred a dated phylogeny for cork and holly oaks using a reduced-representation next-generation sequencing method, restriction site-associated DNA sequencing (RAD-seq), and used D-statistics to investigate gene flow hypotheses. We estimated divergence times using a fossilized birth–death model calibrated with 47 fossils. We used Köppen profiles, selected bioclimatic parameters and forest biomes occupied by modern species toinfer ancestral climatic and biotic niches.

Key Results: East Asian and Western Eurasian cork oaks diverged initially in the Eocene. Subsequently, four Western Eurasian lineages (subsections) differentiated during the Oligocene and Miocene. Evolution of leaf size, form and texture was correlated, in part, with multiple transitions from ancestral humid temperate climates to mediterranean, arid and continental climates. Distantly related but ecologically similar species converged on similar leaf traits in the process.

Conclusions: Originating in temperate (frost-free) biomes, Eocene to Oligocene ranges of the primarily deciduous cork oaks were restricted to higher latitudes (Siberia to north of Paratethys). Members of the evergreen holly oaks (section Ilex) also originated in temperate biomes but migrated southwards and south-westwards into then-(sub)tropical southern China and south-eastern Tibet during the Eocene, then westwards along existing pre-Himalayan mountain ranges. Divergent biogeographical histories and deep-time phylogenetic legacies (in cold and drought tolerance, nutrient storage and fire resistance) thus account for the modern species mosaic of Western Eurasian oak communities, which are composed of oaks belonging to four sections.

In this chapter, we review major classification schemes proposed for oaksby John Claudius Loudon, Anders Sandøe Ørsted, William Trelease, Otto Karl AntonSchwarz, Aimée Antoinette Camus, Yuri Leonárdovich Menitsky, and Kevin C.Nixon. Classifications of oaks (Fig. 2.1) have thus far been based entirely on morphologicalcharacters. They differed profoundly from each other because each taxonomistgave a different weight to distinguishing characters; often characters that arehomoplastic in oaks. With the advent of molecular phylogenetics our view hasconsiderably changed. One of the most profound changes has been the realisation thatthe traditional split between the East Asian subtropical to tropical subgenusCyclobalanopsis and the subgenus Quercus that includes all other oaks is artificial.The traditional concept has been replaced by that of two major clades, each comprisingthree infrageneric groups: a Palearctic-Indomalayan clade including GroupIlex (Ilex oaks), Group Cerris (Cerris oaks) and Group Cyclobalanopsis (cycle-cupoaks), and a predominantly Nearctic clade including Group Protobalanus (intermediateor golden cup oaks), Group Lobatae (red oaks) and Group Quercus (white oaks, with most species in America and some 30 species in Eurasia). In addition, recentphylogenetic studies identified two distinct clades within a wider group of white oaks: the Virentes oaks of North America and a clade with two disjunct endemic species inwestern Eurasia and western North America, Quercus pontica and Q. sadleriana. Themain morphological feature characterising these phylogenetic lineages is pollenmorphology, a character overlooked in traditional classifications. This realisation,along with the now available (molecular-)phylogenetic framework, opens new avenuesfor biogeographic, ecological and evolutionary studies and a re-appraisal of thefossil record. We provide an overview about recent advances in these fields andoutline how the results of these studies contribute to the establishment of a unifyingsystematic scheme of oaks. Ultimately, we propose an updated classification ofQuercus recognising two subgenera with eight sections. This classification considersmorphological traits, molecular-phylogenetic relationships, and the evolutionaryhistory of one of the most important temperate woody plant genera.

The position of Turkey between Europe and Asia makes this region interesting for palaeobotanical investigations. We investigated plant macrofossils from early Miocene deposits of W Turkey (Soma, Manisa) and compiled a catalogue of revised and new plant taxa. We documented 100 fossil-taxa, of which several are new for Turkey (Mahonia aff. pseudosimplex, Ziziphus paradisiaca, Comptonia longirostris, Carya denticulata, Viscum, Fatsia, Pungiphyllum cruciatum). Some previous records are rejected (e.g. Apocynophyllum, Cassia, Castanea, Ficus, Illicium, Liriodendron, Vaccinium). Using modern ecology and taphonomy, we reconstructed palaeoenvironments. We found evidence for a belt of drier, more open habitats with cycads, Dracaena, Mahonia, Smilax miohavanensis, and others. Other vegetation units comprise swamp and riparian forest with few dominants (expressed in highabundance of leaf specimens). On well-drained soils, lowland forest with large-leaved Lauraceae vel Fagaceae and rare elements (Fatsia) was present, while humid temperate broadleaf-deciduous and conifer forest flourished higher up. To infer palaeoclimate we used the Climate Leaf Analysis Multivariate Program (CLAMP) and compared the results with other Miocene localities of Turkey. Early Miocene floras had warm climates (mean annual temperature, MAT, and coldest month temperature, CMMT) with weak precipitation seasonality. MAT and CMMT were cooler during the early middle Miocene. Several fossil-taxa at Soma have biogeographic links with older/coeval localities in C Europe and/or the W Mediterranean region (cycads, Torreya, Dracaena, Smi-lax miohavanensis, Mahonia aff. pseudosimplex, Carya denticulata, Ilex miodipyrena, Pungiphyllum). Few taxa have clear links to C Asian floras (Comptonia longirostris). True E Mediterranean endemics are even rarer (Mahoniaspp., Quercus sosnowskyi).

The early Miocene was a period of major palaeogeographic reorganization in the eastern Mediterranean region, during which time the Anatolian Plateau became subaerial and several intracontinental basins intermittently became connected to the Paratethys and Mediterranean seas. In this paper, we analyse early Miocene vegetation and climate using leaf records, palynological assemblages, and fossil wood at 36 localities from western and central Turkey, most of which have precise age control based on radiometric dating and mammal faunal ages. Using the leaf flora of Güvem (Beş Konak, Keseköy), Climate Leaf-Analysis Multivariate Program (CLAMP) analyses and Köppen signatures were employed to infer a palaeoclimate typical of modern laurel forest regions. Based on the palynological records, abundance of various pollen-taxa was used as a measure of openness of vegetation and regional presence of major tree taxa. Most pollen floras are dominated by tree pollen (ranging from 85 to 98%) and indicated widespread afforestation. In the pollen diagrams, shifts in dominance from swamp forest elements (Taxodioideae) to well-drained forests (Pinaceae) indicate changes in lake levels or phases of basin development. Such shifts may have been associated with the development of more xeric forest vegetation. Wood anatomical features such as false tree rings further may indicate seasonal climate. Pollen diagrams and macrofossils reflect zonal and azonal broadleaf and needleleaf forest and extrazonal open vegetation. The latter occurred in areas with shallow soils on volcanic rocks or limestone (e.g. cycads, Dracaena), or coastal areas (herb dominance). Taxonomic composition and biogeographic affinities suggest laurel forest as a major forest biome on well-drained soils and ecotones between laurel forest and broadleaf deciduous forest biomes. A comparison with younger floras shows that these are neither more diverse nor more warmth-loving despite an increase in global temperature (Mid-Miocene Climatic Optimum) suggesting bottlenecks during previous (Oligocene) cooler times for warmth-loving taxa.

The piggyback basin of Bezhan, southeastern Albania, was formed during the late Neogene and contains Pliocene/Pleistocene deposits. These continental deposits consist of marls, siltstones and clays separated by a thin series of lignite-seams alternating with clays (Bezhan formation). We investigated leaf fossils and dispersed pollen from marls of the upper portion of this formation. Fifty-two plant taxa comprising algae, gymnosperms, and angiosperms were recovered. Of these, at least 19% belong to extant species and less than 16% belong to taxa today extinct in western Eurasia. Tsuga is represented by three pollen taxa with affinities to modern Chinese, Japanese, and North American species. Herbaceous taxa indicative of steppe (Artemisia, Amaranthaceae) occur in low quantities (≤1%)suggesting an interglacial setting. Four vegetation units are recognised: Wet riparian and aquatic vegetation, mesic oak forest, dry sub-Mediterranean woodland, and montane conifer forest. A comparison of the Bezhan flora with well-dated Pliocene and Pleistocene floras of Italy suggests a Calabrian (late early Pleistocene) age for the upper unit. This estimate is based on the abundance of extant taxa, the absence of subtropical taxa, and threshold values of particular taxa (Tsuga, Carya). The findings are in agreement with age estimates for extant tree species from molecular studies.

The early Burdigalian (MN3) plant assemblage of the Güvem area (northwestern Central Anatolia) is preserved in lacustrine sediments of the Dereköy pyroclastics. Its age is well constrained by radiometric dates of basaltic rocks bracketing the pyroclastics, making the Güvem flora one of the extremely few precisely dated early Miocene floras in the Mediterranean region. The rich assemblage of impression fossils comprises ferns and fern allies (2 species), gymnosperms (12 spp.) and angiosperms (129 spp.). Ilex miodipyrena sp. nov. is described as a new fossil-species. The most diverse families in the assemblage are the Fagaceae with 12 taxa and the Fabaceae with 12 leaf morphotypes and one fruit taxon. Aquatic plants are represented by seven taxa, riparian (including palms) and swamp forest elements by >35 taxa, and lianas by three taxa (Smilax spp., Chaneya). The relatively large number of aquatic and riparian/swamp elements is congruent with the rich fish, amphibian and reptile record of the Güvem area. Another characteristic feature of the plant assemblage is the presence of various lobed leaves which show similarities with modern species of different families (e.g. Alangium, various Malvales). Trees and shrubs growing on well-drained soils and forming closed-canopy and open-canopy forests are the most diversified group (>70 taxa). In terms of number of specimens in the collection and based on field observations, by far the most abundant leaf fossils belong to evergreen oaks of Quercus drymeja and Q. mediterranea and to various types of foliage that cannot be assigned to a particular extant or extinct genus of Fagaceae. These sclerophyllous trees must have covered vast areas surrounding the wetlands that developed during the early Miocene in the Güvem Basin. Based on a recent reassessment of the ecology and taxonomic affinity of these trees, they are considered to reflect humid temperate climatic conditions but with a brief drier season during the winter months. These forests are more similar to the laurel forests of the southeastern United States and those stretching in a narrow belt south of the Himalayas to eastern central China. The large number of Fabaceae may indicate the presence of warm subtropical environments but this is difficult to assess, as they are known for having wide ecological ranges today and in the past. All in all, a larger part of the plant taxa point to forested vegetation. This is in agreement with previous palynological studies which detected only small amounts of herbaceous and grass pollen. Open patches of vegetation may have been restricted to river banks and to rocky areas in a volcanic landscape. The biogeographic patterns detected for the early Miocene of the Güvem assemblage are manifold; most taxa are widespread Northern Hemispheric elements. A substantial part of the species migrated from Asia into Europe during the (late) Paleogene and reached Anatolia during the early Miocene (Fagus, Paliurus, Chaneya, Ailanthus, Quercus kubinyii, Davallia haidingeri, Acer angustilobum, A. palaeosaccharinum). Fewer taxa may have been in Anatolia before they migrated to Europe (e.g. Nerium, Smilax miohavanensis, Quercus sosnowskyi). Finally, very few taxa are Anatolian endemics (e.g. Ilex miodipyrena).

Wilf et al. (Research Articles, 7 June 2019, eaaw5139) claim that Castanopsis evolved in the Southern Hemisphere from where it spread to its modern distribution in Southeast Asia. However, extensive paleobotanical records of Antarctica and Australia lack evidence of any Fagaceae, and molecular patterns indicate shared biogeographic histories of Castanopsis, Castanea, Lithocarpus, and Quercus subgenus Cerris, making the southern route unlikely.

 The tree of life is highly reticulate, with the history of population divergence emerging frompopulations of gene phylogenies that reflect histories of introgression, lineage sorting anddivergence. In this study, we investigate global patterns of oak diversity and test the hypothesisthat there are regions of the oak genome that are broadly informative about phylogeny.

 We utilize fossil data and restriction-site associated DNA sequencing (RAD-seq) for 632 individuals representing nearly 250 Quercus species to infer a time-calibrated phylogeny ofthe world’s oaks. We use a reversible-jump Markov chain Monte Carlo method to reconstructshifts in lineage diversification rates, accounting for among-clade sampling biases. We thenmap the > 20 000 RAD-seq loci back to an annotated oak genome and investigate genomicdistribution of introgression and phylogenetic support across the phylogeny.

 Oak lineages have diversified among geographic regions, followed by ecological divergencewithin regions, in the Americas and Eurasia. Roughly 60% of oak diversity traces back to fourclades that experienced increases in net diversification, probably in response to climatic transitionsor ecological opportunity.

 The strong support for the phylogeny contrasts with high genomic heterogeneity in phylogeneticsignal and introgression. Oaks are phylogenomic mosaics, and their diversity may infact depend on the gene flow that shapes the oak genome.

The tree of life is highly reticulate, with the history of population divergence emerging frompopulations of gene phylogenies that reflect histories of introgression, lineage sorting anddivergence. In this study, we investigate global patterns of oak diversity and test the hypothesisthat there are regions of the oak genome that are broadly informative about phylogeny. We utilize fossil data and restriction-site associated DNA sequencing (RAD-seq) for 632individuals representing nearly 250 Quercus species to infer a time-calibrated phylogeny ofthe world’s oaks. We use a reversible-jump Markov chain Monte Carlo method to reconstructshifts in lineage diversification rates, accounting for among-clade sampling biases. We thenmap the > 20 000 RAD-seq loci back to an annotated oak genome and investigate genomicdistribution of introgression and phylogenetic support across the phylogeny. Oak lineages have diversified among geographic regions, followed by ecological divergencewithin regions, in the Americas and Eurasia. Roughly 60% of oak diversity traces back to fourclades that experienced increases in net diversification, probably in response to climatic transitionsor ecological opportunity. The strong support for the phylogeny contrasts with high genomic heterogeneity in phylogeneticsignal and introgression. Oaks are phylogenomic mosaics, and their diversity may infact depend on the gene flow that shapes the oak genome.

The late Early Pleistocene was the last time of equable climate in northern Central Italy, reflected in its large mammal fauna and numerous palynological records. Reliably dated leaf fossils from this time are rare, but provide crucial information onlocal and regional vegetation, biogeographic relationships, and species turnover coinciding with the assembly of modern forest biomes. Here we investigated a rich assemblage of leaf and fruit impressions (63 fossil-taxa) from the Oriolo quarry, Faenza (Ravenna), dated to the latest Calabrian, preserved in transgressive coastal deposits. The fossil assemblage represents riparian vegetation and xeric alluvial habitats in the lowlands and forest vegetation above the alluvial plain. Forest-building trees were deciduous, but comprised several taxa characteristic of open scrubland or forest edges. The composition of the flora reflects a dynamic process of assembling modern forest biomes in western Eurasia. While most taxa correspond to modern submediterranean and temperatewoody species, some others represent late occurrences of taxa today confined to refugia outside Italy (Parrotia, Gleditsia, Pterocarya), and a few are Miocene/Pliocene relics indicating final floristic links with East Asia (Japan) and/or North America (Tsuga cf.chiarugii, Carya cf. minor) and Pleistocene endemism in Italy (Berberis auriolensis, Acer aemilianum).

Sclerophyllous oaks (genus Quercus) play important roles in Neogene ecosystems of south-western Eurasia. Modern analogues (‘nearest living relatives’) for these oaks have been sought among five of six infrageneric lineages of Quercus, distributed across the entire Northern Hemisphere. A revision of leaf fossils from lower Miocene to Pliocene deposits suggests that morphotypes of the Quercus drymeja complex are very similar to a number of extant Himalayan, East Asian, and Southeast Asian species of Quercus Group Ilex and may indicate subtropical, relatively humid conditions. Quercus mediterranea comprises leaf morphotypes that are encountered in modern Mediterranean species of Quercus Group Ilex, but also in Himalayan and East Asian members of this group indicating fully humid or summer-wet conditions. The fossil taxa Quercus drymeja and Q. mediterranea should be treated as morphotype complexes, which possibly comprised different biological species at different times. Quercus mediterranea, although readily recognizable as a distinct morphotype in early to late Miocene plant assemblages, may in fact represent small leaves of the same plants that constitute the Quercus drymeja complex. Based on the available evidence, the taxa [GG1] forming the Q. drymeja complex and Q. mediterranea thrived in fully humid or summer-wet climates. The onset of the modern vegetational context of Mediterranean sclerophyllous oaks is difficult to trace, but may have been during the latest Pliocene/early Pleistocene.

Reconstruction of palaeobiomes, ancient communities that exhibit a physiognomic and functional structure controlled by their environment, depends on proxies from different disciplines. Based on terrestrial mammal fossils, the late Miocene vegetation from China to the eastern Mediterranean and East Africa has been reconstructed as a single cohesive biome with increasingly arid conditions, with modern African savannahs the surviving remnant. Here, we test this reconstruction using plant fossils spanning 14–4 million years ago from sites in Greece, Bulgaria, Turkey, the Tian Shan Mountains and Baode County in China, and East Africa. The western Eurasian sites had a continuous forest cover of deciduous or evergreen angiosperms and gymnosperms, with 15% of 1,602 fossil occurrences representing conifers, which were present at all but one of the sites. Raup–Crick analyses reveal high floristic similarity between coeval eastern Mediterranean and Chinese sites, and low similarity between Eurasian and African sites. The disagreement between plant-based reconstructions, which imply that late Miocene western Eurasia was covered by evergreen needleleaf forests and mixed forests, and mammal-based reconstructions, which imply a savannah biome, throws into doubt the approach of inferring Miocene precipitation and open savannah habitats solely from mammalian dental traits. Organismal communities are constantly changing in their species composition, and neither animal nor plant traits by themselves are sufficient to infer entire ancient biomes. The plant fossil record, however, unambiguously rejects the existence of a cohesive savannah biome from eastern Asia to northeast Africa.

The Gleisdorf Formation (Fm.) deposits in the clay pit at Gratkorn, Styria, Austria, are dated to 12.2–12 Ma,and are of late Middle Miocene age (late Serravallian or Sarmatian). To reconstruct the paleovegetation and estimate the paleoclimate at this important vertebrate site, the palynoflora close to the boundary between the vertebrate-bearing layers of the Gratkorn Fm. and the overlying limnic clay deposits of the Gleisdorf Fm. was investigated. Using the single-grain method, 140 palynomorphs were identified. The palynoflora suggests that the paleovegetation was characterised by well-drained lowland and upland forests, riparian forest, and swamp forests. Depending on the dominating tree species, lowland and upland forests might have had closed or more open canopies. Open habitats included wet meadows and shrublands. In addition, conifers were present in theswampy lowlands and the forested uplands. The most prominent paleoclimatic signatures of the palynoflora indicate a fully humid warm temperate climate, with hot to warm summers and cool winters (Cfa-, Cfb-climate), and a seasonal climate with cool and drier winters and hot to warm and wetter summers (Cwa-, Cwb-climate). Our results align with existing studies bordering the Styrian Basin and support the presence of subtropical to warm-temperate vegetation around Gratkorn during the Sarmatian.

The ‘Coexistence Approach’ is amutual climate range (MCR) technique combinedwith the nearest-living relative (NLR) concept. It has been widely used for palaeoclimate reconstructions based on Eurasian plant fossil assemblages; most of them palynofloras (studied using light microscopy). The results have been surprisingly uniform, typically converging to subtropical, per-humid or monsoonal conditions. Studies based on the Coexistence Approach have had a marked impact in literature, generating over 10,000 citations thus far. However, recent studies have pointed out inherent theoretical and practical problems entangled in the application of this widely used method. But so far little is known how results generated by the Coexistence Approach are affected by subjective errors, data errors, and violations of the basic assumptions. The majority of Coexistence Approach studies make use of the Palaeoflora database (the combination of which will be abbreviated to CA + PF). Testing results produced by CA + PF studies has been hindered by the general unavailability of the contents in the underlying Palaeoflora database; two exceptions are the mean-annual temperature tolerances and lists of assigned associations between fossils and nearest-living relatives. Using a recently published study on the Eocene of China,which provides the first and only insight into the data structure of the Palaeoflora database,we compare the theory and practice of Coexistence Approach using the Palaeoflora database (CA+PF).We show that CA+PF is riddled by association and climate data error.We reveal flaws in the application of the Coexistence Approach,which is often in stark contrast to the theory of the method. We show that CA + PF is highly vulnerable against numerous sources of errors, mainly because it lacks safeguards that could identify unreliable data. We demonstrate that the CA+PF produces coherent, pseudo-precise results even for artificially generated, randomplant assemblages. AlternativeMCR-NLR methods can surpass the most imminent deficits of the Coexistence Approach, and may be used as a stop-gap until more accurate bioclimatic and distribution data on potential Eurasian NLRs, and theoretically and statistically robust methods will become available. Finally, general guidelines are provided for the future application of methods using the mutual climatic range with nearest living relatives approach when reconstructing climate from plant fossil assemblages.

This chapter provides a review of Cenozoic plant assemblages from the sub-arctic North Atlantic region and their biogeographic implications. Previous work is reviewed and new data are presented that considerably change our understanding of the role of the northern North Atlantic for plant dispersal and evolution of plant lineages during the Paleogene and Neogene. Paleogene plant fossils in this region are known from West and East Greenland, the Faroe Islands and Scotland. In contrast to the widely held view that most Paleogene plant taxa of Greenland belong to extinct lineages, we provide evidence for the presence of several extant genera in these floras (e.g. Fagus, Quercus). Thus, Engler’s hypothesis about the ‘Arcto-Tertiary element’ remains a fundamental hypothesis about the origin of northern temperate tree genera. In general, a remarkable diversity of extinct and modern lineages of Fagaceae is documented for Palaeocene and Eocene floras. Neogene fossils are found in Iceland and provide records of climate evolution in the sub-arctic North Atlantic and of the duration of a functioning land bridge for plant migration between North America and Europe. Counter to the traditional view suggesting a functioning land bridge only during the Paleogene, there isnow convincing evidence that this link was available for plants until the latest Miocene. This has important implications for understanding low genetic differentiation documented in extant plant groups having a disjunct distribution in northern temperate regions of Europe, North America and East Asia. Relatively warm conditions persisted in the sub-arctic North Atlantic until the end of the Zanclean (early Pliocene) based on plant fossil evidence from Iceland. The shift to modern tundra conditions occurred during the Piacenzian (late Pliocene) and is documented in the Pliocene and Pleistocene fossil plant assemblages of Iceland.

Plant macrofossils from the lignite mines of Eskihisar, Tınaz, and Salihpaşalar (Yatağan Basin, southwestern Anatolia) were investigated. The fossils were collected from marls overlying the exploited lignite seams and represent three subbasins within the main Yatağan Basin. The age of the Eskihisar lignite seam is well constrained by vertebrate fossils (MN 6, middle Miocene). Further, lithological and palynological correlation suggests that the lignite seams and overlying marls in the three lignite mines were formed at the same time. Three distinct zonal vegetation types are reflected in the local plant assemblages: (i) In Eskihisar, Fagus and evergreen Quercus mediterranea-Q. sosnowskyi communities formed important parts of the zonal vegetation along with the deciduous Quercus kubinyii; (ii) in Tınaz, Quercus sosnowskyi-Q. mediterranea-Q. drymeja communities occurred, while Fagus is rarely encountered in the macrofossil record. (iii) In Salihpaşalar, Quercus mediterranea and Q. drymeja are the most abundant elements, while Fagus and Q. sosnowskyi are absent or nearly so. This demonstrates that local environmental conditions within a geographically restricted region varied and probably were controlled by slope aspects, edaphic conditions, and river drainage. Overall, the zonal vegetation is characterized by a high diversity of evergreen and deciduous oaks belonging to Quercus subgen. Cerris sect. Ilex and sect. Cerris and the local dominance of Fagus. The riparian vegetation was dominated by Populus, Salix and Acer, whereas Alnus and taxodiaceous Cupressaceae and ferns were very rare or absent. The mass occurrence of Quercus sosnowskyi in the Yatağan Basin floras is biogeographically interesting, as this distinct sclerophyllous oak has previously been known to occur only in late Miocene sediments of northern Greece and Abkhasia (Georgia).

Plant macro fossils from the lignite mines of Eskihisar, Tınaz, and Salihpaşalar (Yatağan Basin, southwestern Anatolia) were investigated. The fossils were collected from marls overlying the exploited lignite seams and represent three subbasins within the main Yatağan Basin. The age of the Eskihisar lignite seam is well constrained by vertebrate fossils (MN 6, middle Miocene). Further, lithological and palynological correlation suggests that the lignite seams and overlying marls in the three lignite mines were formed at the same time. Three distinct zonal vegetation types are reflected in the local plant assemblages: (i) In Eskihisar, Fagus and evergreen Quercus mediterranea-Q. sosnowskyi communities formed important parts of the zonal vegetation along with the deciduous Quercus kubinyii; (ii) in Tınaz, Quercus sosnowskyi-Q. mediterranea-Q. drymeja communities occurred, while Fagus is rarely encountered in the macrofossil record. (iii) In Salihpaşalar, Quercus mediterranea and Q. drymeja are the most abundant elements, while Fagus and Q. sosnowskyi are absent or nearly so. This demonstrates that local environmental conditions within a geographically restricted region varied and probably were controlled by slope aspects, edaphic conditions, and river drainage. Overall, the zonal vegetation is characterized by a high diversity of evergreen and deciduous oaks belonging to Quercus subgen. Cerris sect. Ilex and sect, Cerris and the local dominance of Fagus. The riparian vegetation was dominated by Populus, Salix and Acer, whereas Alnus and taxodiaceous Cupressaceae and ferns were very rare or absent. The mass occurrence of Quercus sosnowskyi in the Yatağan Basin floras is biogeographically interesting, as this distinct sclerophyllous oak has previously been known to occur only in late Miocene sediments of northern Greece and Abkhasia (Georgia). 

We report a new type of fossil margin galls arranged in a linear series on dicot leaf impressions from the latest Neogene (Pliocene) sediments of the Chotanagpur Plateau, Jharkhand, eastern India. We collected ca. 1500 impression and compression leaf fossils, of which 1080 samples bear arthropod damage referable to 37 different damage types (DT) in the ‘Guide to Insect (and Other) Damage Types in Compressed Plant Fossils’. A few leaf samples identified as Ipomoea L. (Convolvulaceae) have specific margin galls that do not match any galling DT previously described. This type of galling is characterized by small, linearly arranged, irregular, sessile, sub-globose, solitary, indehiscent, solid pouch-galls with irregular ostioles. The probable damage inducers of the present galling of the foliar margin might be members of Eriophyidae (Acari). The new type of gall suggests that marginal gall-inducing mites on leaves of Ipomoea did not change their host preference at the genus level since the Pliocene. The development of marginal leaf galling in Ipomoea is linked to extrafloral nectaries that do not offer protection against arthropod galling but indirectly protect the plant against herbivory from large mammals.

The Pliocene flora of Frankfurt am Main described by Karl Mädler during the first half of the twentieth century is a key flora for the European Pliocene. In the present study, we revised the leaf fossil taxa described by Mädler and investigated plant material collected after Mädler’s publication. The revised and augmented floral list comprises seven new species and some new combinations of taxa described by Mädler. In total, 16 gymnosperm species in 15 genera and 73 angiosperm species (of which 15 could not be assigned to a genus) in 40 genera are recognised in the leaf record. Main characteristics of the flora are the high diversity of conifers, the diverse assemblage of exclusively deciduous Fagaceae, including six species of oaks, and the high diversity of Rosaceae. These features indicate cool temperate climatic conditions (comparable to Lugano in southern Switzerland). Angiosperm genera that are today confined to North America and/or East Asia (Eucommia, Magnolia and Sassafras) also are deciduous, whereas evergreen taxa are shrubs typical of the understorey (Buxus, Ilex, Pachysandra, Prunus lusitanica type) and Viscum. Eighteen taxa recorded in the Pliocene of Frankfurt am Main are today absent from western Eurasia and eastern North America, and 25 taxa are absent from western North America. This shows (i) a strong biogeographic link of the Pliocene flora of Frankfurt am Main with East Asia, (ii) surprisingly high levels of speciation (Pliocene endemisms) and (iii) that the European flora was more diverse in woody species shortly before the onset of major Pleistocene glaciations than today.

The Pliocene flora of Frankfurt amMain described by Karl Mädler during the first half of the twentieth century is a key flora forthe European Pliocene. In the present study, we revised the leaf fossil taxa described by Mädler and investigated plant materialcollected after Mädler’s publication. The revised and augmented floral list comprises seven new species and some new combinationsof taxa described by Mädler. In total, 16 gymnosperm species in 15 genera and 73 angiospermspecies (of which 15 couldnot be assigned to a genus) in 40 genera are recognised in the leaf record.Main characteristics of the flora are the high diversity ofconifers, the diverse assemblage of exclusively deciduous Fagaceae, including six species of oaks, and the high diversity ofRosaceae. These features indicate cool temperate climatic conditions (comparable to Lugano in southern Switzerland).Angiosperm genera that are today confined to North America and/or East Asia (Eucommia, Magnolia and Sassafras) also aredeciduous, whereas evergreen taxa are shrubs typical of the understorey (Buxus, Ilex, Pachysandra, Prunus lusitanica type) andViscum. Eighteen taxa recorded in the Pliocene of Frankfurt am Main are today absent from western Eurasia and eastern NorthAmerica, and 25 taxa are absent from western North America. This shows (i) a strong biogeographic link of the Pliocene flora ofFrankfurt amMain with East Asia, (ii) surprisingly high levels of speciation (Pliocene endemisms) and (iii) that the Europeanflora was more diverse in woody species shortly before the onset of major Pleistocene glaciations than today.

The Late Oligocene to Early Miocene flora of the Ban Pa Kha Subbasin (Li Basin, northern Thailand) provides a record of montane dry tropical oak-pine forests. The rich ensemble of Fagaceae typical of these forests might have existed in the wider region of Southeast Asia since Eocene times and various fossil plant assemblages represented both lowland (Fagaceae, Dipterocarpaceae) and upland (Fagaceae, Pinaceae) tropical forests. These findings are in conflict with previous interpretations of vegetation development in northern Thailand, which stressed that stratigraphically older (possibly Late Oligocene) spore and pollen assemblages in northern Thailand were markedly different from the modern tropical flora and had a distinct northern temperate character. A major change in climate would have caused a dramatic shift to tropical conditions since the Mid-Miocene. Considering palaeobotanical data from adjacent regions in Southeast Asia, we suggest that differences in spore and pollen assemblages in intermontane basins in northern Thailand are more likely to represent different facies and lowland/upland settings. Assembly of these forest ecosystems, typically comprising Quercus sections Cyclobalanopsis, Ilex and Quercus, pollen of Castaneoideae with affinities to Castanopsis and Lithocarpus, and extinct fagaceous genera such as Eotrigonobalanus, most probably dates back to the Eocene. The absence of oaks of Quercus section Cerris in the spore and pollen assemblage of the Ban Pa Kha Subbasin, despite this group being part of the modern vegetation, might reflect the late arrival (secondary radiation) of this chiefly temperate group in tropical Southeast Asia.

Measuring biological diversity is a crucial but difficult undertaking, as exemplified in oaks where complex patterns of morphological, ecological, biogeographical and genetic differentiation collide with traditional taxonomy, which measures biodiversity in number of species (or higher taxa). In this pilot study, we generated high-throughput sequencing amplicon data of the intergenic spacer of the 5S nuclear ribosomal DNA cistron (5S-IGS) in oaks, using six mock samples that differ in geographical origin, species composition and pool complexity. The potential of the marker for automated genotaxonomy applications was assessed using a reference data set of 1,770 5S-IGS cloned sequences, covering the entire taxonomic breadth and distribution range of western Eurasian Quercus, and applying similarity (blast) and evolutionary approaches (maximum-likelihood trees and Evolutionary Placement Algorithm). Both methods performed equally well, allowing correct identification of species in sections Ilex and Cerris in the pure and mixed samples, and main lineages shared by species of sect. Quercus. Application of different cut-off thresholds revealed that medium- to high-abundance (>10 or 25) sequences suffice for a net species identification of samples containing one or a few individuals. Lower thresholds identify phylogenetic correspondence with all target species in highly mixed samples (analogous to environmental bulk samples) and include rare variants pointing towards reticulation, incomplete lineage sorting, pseudogenic 5S units and in situ (natural) contamination. Our pipeline is highly promising for future assessments of intraspecific and interpopulation diversity, and of the genetic resources of natural ecosystems, which are fundamental to empower fast and solid biodiversity conservation programmes worldwide.

The partial female inflorescence reported here provides an important addition to acorns of Castanopsis described from middle Eocene strata of Europe. Furthermore, the intercontinental distribution of Castanopsis in the Eocene is confirmed. The amber fossil also broadens the picture of the Baltic amber source area, indicating oligotrophic, sandy, bog-like habitats. Finally, this study underscores the great benefit of SRμCT as a powerful tool to investigate plant inclusions from amber in a nondestructive way.

Eocene Baltic amber forms the largest amber deposit worldwide; however, its source vegetation and climateare much debated. Representatives of the oak family (Fagaceae) were abundant in the Baltic amber source areabased on numerous inclusions of staminate inflorescences or individual florets, previously assigned to Castanea andQuercus. However, the actual generic and infrageneric diversity of Fagaceae from Baltic amber remained unknown.Using flower characteristics and section-diagnostic in situ pollen of staminate inflorescences and detached floret inclusions,we describe 18 fossil-species of Fagaceae making this family by far the most diverse plant family preservedin Baltic amber. We substantiate the occurrence of the Castaneoideae, Quercoideae (Quercus sect. Cyclobalanopsis/ Lobatae; Q. sect. Lobatae; Q. sect. Protobalanus), Trigonobalanoideae and the extinct genus Eotrigonobalanus.Among the 18 fossil-species, six are described as new: Q. aimeeana, Q. casparyi, Q. multipilosa, E. campanulata,E. conwentzii, E. longianthera; and one new combination is published: Q. brachyandra (≡ Castanea brachyandra).In addition, a lectotype is designated for the name Quercites meyerianus and neotypes are designated for the namesCastanea inclusa and Quercus longistaminea (≡ C. longistaminea). Members of the Fagaceae probably inhabitedazonal and zonal vegetation types of the amber source area, including bottomland flood-plains and stream banks(Q. sect. Lobatae), dry habitats (Q. sect. Lobatae, Q. sect. Protobalanus), peaty soils, riparian and swamp forests(Castanopsis, Eotrigonobalanus), as well as mixed mesophytic forests (castaneoids, Quercoideae, trigonobalanoids).Affinities to extant North American and E to SE Asian floras support the recent notion that late Eocene Baltic amber(38 – 34 Ma) was formed in a warm-temperate climate.