Combined LM and SEM study of the middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin, Austria: Part V. Magnoliophyta 3 – Myrtales to Ericales

Abstract The continued investigation of the middle Miocene palynoflora from the Lavanttal Basin reveals numerous additional angiosperm taxa. The Myrtales to Ericales pollen record documented here comprises 46 different taxa belonging to Onagraceae (Ludwigia), Ericaceae (Craigia, Reevesia, Tilia), Anacardiaceae (Pistacia), Rutaceae (Zanthoxylum), Sapindaceae (Acer), Santalaceae (Arceuthobium), Amaranthaceae, Caryophyllaceae, Polygonaceae (Persicaria, Rumex), Cornaceae (Alangium, Cornus, Nyssa), Ebenaceae (Diospyros), Ericaceae (Andromeda, Arbutus, Empetrum, Erica), Sapotaceae (Pouteria, Sideroxylon), Styracaceae (Rehderodendron) and Symplocaceae (Symplocos). Köppen signatures of potential modern analogues of the additional fossil woody elements confirm the hypothesis of a subtropical (Cfa, Cwa) climate at lower elevations and subsequent transition into a temperate climate with altitudinal succession (Cfa → Cfb/Dfa → Dfb; Cwa → Cwb → Dwb-climate). The fossil plants represent different vegetation units, from wetland lowlands to well-drained montane forests. Many of the fossil taxa have potential modern analogues that can be classified as nemoral and/or meridio-nemoral and/or semihumid-meridional vegetation elements. New is the recognition of oreotropical elements, which are direct indicators for a substantial altitudinal gradient.

accounting for 21 of the here recorded taxa. The asterids are represented by two orders, the Cornales and Ericales, in total, 25 taxa. Based on the potential modern analogues of the fossils we can confirm and refine our preliminary hypotheses about palaeovegetation and palaeoecology.

Material and methods
For a detailed account on the geographical position, geology and age of the Lavanttal Basin and its surroundings, sedimentology and palaeoenvironment, preservation of organic matter, as well as detailed information on the sediment samples and preparation methods see .
The fossil pollen grains were investigated both by light microscopy (LM) and scanning electron microscopy (SEM) using the single-grain method as described by Zetter (1989) and Halbritter et al. (2018). Pollen micrographs in compiled figures show the same single pollen grain photographed with LM (sometimes rotated and/or at different focus level) and with SEM (overview and close-up of sculpture).
We used 'Köppen signatures' (Denk et al. 2013) to summarise the climatic niche occupied by potential modern analogues (species groups, genera) of the determined pollen taxa. We also categorised the potential modern analogues as climate-dependent vegetation elements adapting the concepts of Schroeder (1998) to provide a more generalised account of the palaeovegetation in relation to modern-day vegetation zones (cf. Denk et al. 2013, figure 2;Grímsson et al. 2016, figure 2). Here, we recognise two additional climate-vegetation categories, the 'oreotropic' (Figure 1) and 'austral' elements (Schroeder 1998) to accommodate species with hygric and thermic preferences not covered by the data set used by Denk et al. (2013) and Grímsson et al. (2016). Oreotropic species (Schröder 1998) are found in fully humid temperate to winter-dry temperate climates ([A], Cfa, Cfb, Cwa, Cwb; sensu Köppen-Geiger in Kottek et al. 2006) along altitudinal thermic successions in low latitudes characterised by tropical climates. Climatically they are hence similar to meridio-nemoral elements, but are exclusively found in mountainous areas within the tropics and, in contrast to meridio-nemoral elements, show no physical latitudinal connection to the nemoral or boreal vegetation zones. Austral species are adapted to thermic and humidity conditions of the Southern Hemisphere similar to the meridional and nemoral zones of the Northern Hemisphere including temperate fully humid to summer-and winter-dry climates with hot to warm summers (Cfa, Cfb, Csa, Csb, Cwa, Cwb; sensu Köppen-Geiger in Kottek et al. 2006). The climate data for all potential modern analogues of the fossil taxa presented herein are listed in Supplementary File S1.
Note that we use here two, semantically partly overlapping concepts that have caused much confusion in palaeoclimatic and palaeovegetational literature when not strictly applied. For the general climate/vegetation bands/zones, we make use of the latitudinal-based modification of Köppens climate classification by Trewartha (1954;Trewartha & Horn 1980), which recognises a subtropical zone (addressed also as 'subtropics') between the equatorial tropical zone ('tropics') and the (fully or 'cool') temperate zone at mid-latitudes (c. 30°-60°). The Köppen signatures refer strictly to Köppen's system that has no explicit subtropical zone. Thus, when we speak of 'warm temperate climates' (Köppen's C-climates), this includes climates typical for the (low-to mid-altitude) subtropics with hot and/ or dry summers (Cfa, Cwa, Csa, Csb p.p.) but also climates typical for the warmer part of the temperate zone (Cfb, Cwb, rest of Csb) and climates that, according Trevartha's system, transition into the boreal (also known as 'cold temperate') zone (Cfc, Cwc). Vegetation-wise, Köppen's original system is more applicable since many dominant and common taxa of the subtropics extend into the warmer part or deep into the temperate zone. The transition from the subtropical into the temperate zone is vegetation-wise hardly visible in North America or East Asia due to the absence of major east-west physical barriers.

Systematic palaeontology
All descriptions of angiosperm pollen presented herein include the most diagnostic features observed both in LM and SEM. The pollen terminology follows mostly Punt et al. (2007, LM) and Halbritter et al. (2018, SEM). The classification of orders and families follow APG III (2009). Families and genera are arranged in alphabetical order. Remarks. -The pollen morphology of Onagraceae has been studied in detail using LM, SEM and transmission electron microscopy (TEM) by Ting (1966), Brown (1967), Skvarla et al. ( , 1978, Praglowski et al. (1983Praglowski et al. ( , 1987Praglowski et al. ( , 1988Praglowski et al. ( , 1994, Patel et al. (1984), Keri and Zetter (1992), Punt et al. (2003), and Makbul et al. (2008). These studies show that Onagraceae produce distinct pollen types that cannot be confused with pollen from any other angiosperm family (cf. Patel et al. 1984), making it also possible to identify fossil Onagraceae pollen/ tetrads at generic level.
Ecological implications. -Ludwigia displays a cosmopolitan, but mainly pantropical, distribution with some of its 82 species occurring on every continent except Antarctica. These are mostly small herbaceous annual or perennial plants (Wagner et al. 2007). Most extant Ludwigia species are water plants. It is very likely that both Ludwigia sp. 1 and Ludwigia sp. 2 represent small herbaceous plants that grew in lakes or along their shorelines in the lowland wetlands of the Lavanttal Basin.
Remarks. -Pollen morphology of Craigia yunnanensis W.W. Smith et W.E. Evans has been documented using LM and SEM by Long et al. (1985) and Kvaček et al. (2002). In LM the apertures of Craigia pollen is characterised by a circular horseshoe-like nexine thickening best observed in optical cross-section (Kvaček et al. 2002, plate V). In Tilia, this thickening is much broader/wider and less convex (Perveen et al. 2004, figure 4). Tilia pollen is also larger than the pollen of Craigia, and the sculpture of Tilia pollen observed with LM is more prominent.
Fossil record. -The macrofossil record of Craigia, including leaves and fruits, is summarised in Kvaček et al. (2005). The record shows that this genus had a much wider distribution during the Cainozoic, with numerous fossils from the early Oligocene to late Pliocene of Western Eurasia, late Eocene/Oligocene of Spitsbergen, the Paleocene to Miocene of East and Central Asia, and the middle Eocene to middle Miocene of North America (e.g. Kvaček 2004;Kvaček et al. 2005;Jin et al. 2009;Liu et al. 2012). Fossil Craigia pollen is also well represented in the palynological record of the Cainozoic (e.g. Kvaček et al. 2002;Zetter et al. 2002), but the grains have mostly been lumped into various fossil species (along with Tilia type pollen) under the form-genus Intratriporopollenites Pflug et Thomson (cf. Stuchlik et al. 2014). even be extinct. Craigia yunnanensis are small deciduous trees, up to 20 m tall, occurring mostly in open forests, at an elevation of 500 to 1600 m, in southern China and adjacent northernmost Vietnam (Ya et al. 2007b). Today's Craigia prefer warm temperate climates with dry winters and hot summers (Cwa; semihumid-meridional vegetation element; File S1), with mean annual temperatures (MATs) ranging from 13 to 21°C, annual precipitation of 1074 to 1688 mm, and coldest mean month temperature of 6.3 to 14.2°C (Fang et al. 2009). According to Kvaček et al. (2005), C. yunnanensis is usually confined to mesic notophyllous evergreen broadleaved forests, with some populations occurring at higher elevations in mixed conifer and evergreen broadleaved forests or mixed mesophytic forests. Since macrofossils of Craigia (e.g. C. bronnii, Dombeyopsis lobata) are often associated with plants of moist habitats, Kvaček (2004) suggested that during the Miocene the Central European Craigia were part of wetland forest vegetation thriving in backswamp forests and along streams and in deltas. Based on the Lavanttal plant assemblage such a scenario is possible; taking into account its present habitat, however, it cannot be ruled out that Craigia was only/also part of more well-drained forests at higher elevation surrounding the Lavanttal Basin.
Fossil record. -Reevesia pollen is morphologically unique and easy to identify. For this reason, even though it occurs only in low numbers in palynological samples, it has a well-established fossil record. The fossil pollen record of Reevesia and Reevesiapollis Krutzsch has been summarised in detail by Krutzsch (1970b), Petrov and Drazheva-Stamatova (1972), Raine et al. (2011), andStuchlik et al. (2014). These accounts suggest that Reevesia had a wide continental European distribution, with records extending from the middle Paleocene until the Pliocene, but with a peak occurrence during the Miocene. Pollen of this genus is also known from the Eocene to Miocene of Russia and from the Miocene to Pleistocene of New Zealand.
Ecological implications. -The genus Reevesia comprises about 20 species of evergreen and deciduous trees (8-18 m tall) with a South Asian distribution (excluding the two Central American species alternatively placed in Veeresia). Fifteen of the species occur in China, where 12 of them are endemic (Ya et al. 2007a). Most of the Chinese Reevesia occur in warm temperate climate with dry winter and hot summer (Cwa), a third of the taxa occur (also) in equatorial savannah climate with dry winter (Aw-climate), and one third (also) in fully humid warm temperate climate with hot summer (Cfa), in combination, the genus can be characterised as a tropical-meridional element (File S1). The majority of the taxa endure a MAT in the range of 14 to 25°C, and an annual precipitation somewhere between 1268 and 2992 mm (Fang et al. 2009). All the Reevesia species in China have very restricted distributions except for R. pubescens (Aw, Cwa, Cfa); it grows under a MAT of 2 to 25°C, and annual precipitation of 741 to 2435 mm (Fang et al. 2009). Reevesia pubescens Mast. is the only species extending into regions with winter frost, enduring coldest mean month temperatures down to −7°C (Fang et al. 2009). According to Ya et al. (2007a) Reevesia occurs in both open valley forests as well as dense montane forests; it can also be found on forested slopes/hillsides and along riverbanks. Based on habitats of the potential modern analogues of the Lavanttal fossils, especially R. pubescens, the Miocene Reevesia was likely part of forest vegetation surrounding the basin, occurring in well-drained valley-and hillside forests, above the main wetlands.
Genus Tilia L.
Tilia sp. 1 ( Figure 4G- Grímsson et al. 2017b) and unknown or extinct groups (Gastaldo et al. 1998). True Tilia pollen is not known from western Eurasian sediments prior to the Oligocene, but is frequent in Miocene sediments and has been documented, among others, from Iceland (Denk et al. 2011), Germany (Ferguson et al. 1998), Poland (Stuchlik et al. 2014), Austria (Kovar-Eder et al. 1998;Zetter 1998) and Turkey (Bouchal et al. 2016b(Bouchal et al. , 2017Bouchal 2019 Jones 1968;Ya et al. 2007b;Pigott 2012;Hanes 2015). Tilia americana L. covers a wide geographical region in eastern North America, occurring in both fully humid warm temperate climates and snow climates, with either hot or warm summers (Cfa, Cfb, Dfa, Dfb; nemoral vegetation element; File S1). In China, most Tilia species occur in warm temperate climates with hot summers that are either fully humid (Cfa) or winter-dry (Cwa). Only some East Asian Tilia species (≤ 6 spp.) extend into cooler areas with warm summers (Cwb) or (≤ 4 spp.) extend into snow climates (Dwa, Dwb). None of the Tilia species in China are particularly prominent in lowland wetland vegetation, but are predominantly reported from altitudes above 600 m and even occurring in mountain forests up to c. 4000 m (Ya et al. 2007b).
Only very few of the Tilia species (≤ 3 spp.) in China extend into areas where the MAT drops below 0°C, most of the taxa (≥ 11 spp.) occur in areas where the MAT is above 5°C, and the annual precipitation ranges between 250 and 1997 mm (Fang et al. 2009). According to Ya et al. (2007b) Tilia trees in China are components of evergreen or mixed evergreen and deciduous forests, occurring on welldrained steep valley slopes and mountainsides. A similar habitat is suggested for the middle Miocene Tilia from the Lavanttal Basin.
Fossil record. -The European macrofossil record of Pistacia is meagre. According to Mai (1995b), leaf and fruit records are rare and confined to the middle Oligocene and Pliocene of Italy, and the Miocene of Germany, Hungary and Austria. Pistacia pollen is also very rare. So far, the only European records are from the Miocene of France and Spain (cf. Muller 1981).
Ecological implications. -The recently revised classification of Pistacia by Al-Saghir and Porter (2012) recognised nine species (and five subspecies). Pistacia are deciduous or evergreen shrubs to small trees (up to 20 m tall). The genus has a disjunct northern hemispheric distribution, with a single species in south-western United States and Central America (P. mexicana Humb.), five species in Mediterranean Europe and Africa (P. atlantica Desf., P. chinensis Bunge, P. terebinthus L., P. vera L., P. lentiscus L.), six species in West to Central Asia (P. atlantica, P. chinensis, P. eurycarpa Yalt., P. khinjuk Stocks, P. vera, P. lentiscus), and two species in East Asia (P. chinensis, P. weinmannifolia J. Poiss. ex Franch.) (Al-Saghir & Porter 2012). The current diversification centre of Pistacia, the Mediterranean region (summer-dry warm temperate climates), is believed to have been established following the latest Miocene (Xie et al. 2014). Hence, the Mediterranean (and alike) Pistacia species cannot be used as informative modern-day analogues for the Lavanttal fossils. This leaves the East Asian species as potential modern analogues of the fossil taxon, P. chinensis and P. weinmannifolia. The former shows a disjunct distribution across Asia into Africa. In China, P. chinensis has a wide distribution, thriving in winter-dry equatorial savannah climates (Aw) to fully humid warm temperate climate with hot summers (Cfa), and enduring coldest mean month temperatures down to −13°C (Fang et al. 2009). It occurs alongside P. weinmannifolia in warm temperate climates with hot or warm summer (Cwa, Cwb;), hence, the genus can be classified as a tropical-meridional vegetation element (File S1). Both taxa are found in hill and mountain forests, growing mostly on hard substrates (rocky soils and limestone). Pistacia chinensis occurs at an elevation between 100 and 3600 m, and P. weinmanniifolia at elevation between 500 and 2700 m (Tianlu & Barfod 2008). The Miocene Pistacia from Lavanttal was most likely growing outside the lowland wetland area, on well-drained hills and mountain slopes, especially on hard or rocky substrate.
Remarks. -Pollen of extant Zanthoxylum has been studied in LM by Huang (1972), Barth (1980), Wang et al. (1995), and Mayer (1996), in SEM by Barth (1980), Mayer (1996), Li et al. (2011), andCao et al. (2014), and in TEM by Barth (1980). The pollen morphology of Zanthoxylum is unique Miocene palynoflora from Austria within Rutaceae (Mayer 1996). The suite of characters that permit identification of this genus include long and bow-like colpi and lalongate rectangular endopori observed with LM. Also, margins of endopori perpendicular to polar axis are thickened (LM), and the sculpture is striato-reticluate to striate. The only genus producing similar pollen is Toddalia, but their pollen is smaller, spheroidal (versus prolate in Zanthoxylum), and the colpi are shorter.
Fossil record. -According to Gregor (1989) and Mai (1995b) Zanthoxylum has a reliable fossil seed record from the middle Eocene to Pliocene in Europe, the Oligocene to Pleistocene of Asia, and the Miocene of North America. The leaf record is scarce and doubtful (Mai 1995b). This pollen type has rarely been reported (e.g. Gastaldo et al. 1998;Ferguson et al. 1998)  Remarks. -Even though the two Zanthoxylum pollen types are very similar in size and outline they unambiguously belong to two different species because Zanthoxylum sp. 2 has much narrower striae (0.2-0.3 versus 0.6-0.7 µm wide) and is also striatoreticulate over most of the pollen surface versus striate in Zanthoxylum sp. 1.
Fossil record. -The massive macrofossil record (leaves and samaras) of Acer has been summarised by numerous authors including Walther (1972), Tanai (1983), Wolfe and Tanai (1987), Oterdoom (1994), Mai (1995b), Boulter et al. (1996), Manchester (1999), McClain (2000), and Grimm et al. (2007). Based on these accounts, the current consensus is that the earliest accepted Acer fossils are from the Paleocene of North America. Acer is then believed to have dispersed across Beringia into Asia during the Eocene and finally reaching Europe during the Oligocene. Acer was one of the most speciesrich and widely distributed woody genera in the Miocene of Europe (e.g. Walther 1972;Mai 1995b;Boulter et al. 1996). In western Eurasia, striate Acer pollen grains, similar to the fossils, have been documented among others from the Oligocene and Miocene of Germany (Schmid 2000;Kottik 2002), and the Miocene of Iceland (Denk et al. 2011), Poland (Stuchlik et al. 2014), and Turkey (Bouchal et al. 2016b(Bouchal et al. , 2017Bouchal 2019;Denk et al. 2019 palmatum Thunb. (section Palmata), the latter cultivated worldwide as a garden ornamental. The striate fossil Acer sp. 1 and Acer sp. 2 pollen types correspond to pollen from numerous modern North American and Eurasian taxa and it is, therefore, difficult to affiliate them to any of the modern, genetically supported sections. These pollen grains could have originated from lowland wetland trees or from individuals that were part of well-drained highland or even mountain forests surrounding the basin.
Ecological implications. -Acer negundo are deciduous trees or shrubs. The species has a vast range throughout North America (Canada, United States,

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F. Grímsson et al. Mexico, and Guatemala), occurring mostly in forested valleys, along riverbanks, and in mixed riparian forests (van Gelderen 1994). Concerning its vast distribution A. negundo has a wide climate range, occurring under various warm temperate and snow climates (Cfa, Cfb, Dfa, Dfb, Cwa, Dwa, Csb; predominantly a nemoral vegetation element; File S1). Acer saccharinum are deciduous trees (up to 40 m tall) native to eastern and central North America (eastern United States, Canada) (van Gelderen 1994). They occur mostly in lowland wetlands and are found in river floodplains, along streams and lakes, and even in swamps (Gabriel 1990b). They grow under fully humid warm temperate to snow climates with hot or warm summers (Cfa, Cfb, Dfa, Dfb; nemoral vegetation element; File S1). It is possible that the plants producing the Acer sp. 3 pollen grains were European counterparts of the North American A. negundo or A. rubrum-saccharinum lineages and part of the lowland wetland vegetation during the Miocene of Lavanttal, occurring in mixed forest along lakes and streams, and floodplains and backswamps.
Remarks. -This pollen type is much larger than the Acer sp. 1-3. Its sculpture type is different to the long striae of Acer sp. 1 and the radially branching striae observed in Acer sp. 3. This pollen type falls within the striato-reticulate (or microreticulate) group of Biesboer (1975). Modern taxon known to produce similar pollen is A. saccharum, a widespread, morphologically variable species that now includes all North American members of sect. Acer (Adams & Morton 1976;van Gelderen 1994).
Ecological implications. -Acer saccharum are deciduous trees, up to 40 m tall, occurring in eastern Canada and central and eastern United States to Mexico and Guatemala (van Gelderen 1994). Acer saccharum is not a wetland taxon and does not occur in swamps but prefers well-drained slopes at intermediate elevation. It is found in rich forests, on slopes, in ravines and valleys, and near streams. In the Mixed Mesophytic Climax Forest of the eastern United States, A. saccharum appears as a dominant member of the forest canopy in association with Aesculus, Fagus, Liriodendron, Quercus, Tilia and Tsuga (Godman et al. 1990;Gabriel 1990a). Acer saccharum is growing under fully humid warm temperate to snow climates with hot or warm summers (Cfa, Cfb, Dfa, Dfb; nemoral vegetation element; File S1). It is possible that the trees producing this type of Acer pollen, during the Miocene of Lavanttal, were found at moderate elevation above the main wetland as part of ravine vegetation and welldrained species-rich hillside and slope forests. Remarks. -Pollen morphology of the genus was investigated using LM and SEM by Hawksworth and Wiens (1972), also listing all previous studies on the pollen morphology of Arceuthobium. Fossil pollen affiliated to modern Arceuthobium has been assigned to the pollen form-genus Spinulaepollis Krutzsch (e.g. Krutzsch 1962;Stuchlik et al. 2014).
Fossil record. -The pre-Quaternary fossil record of Arceuthobium is scarce and mostly based on dispersed pollen. The earliest pollen records are from the middle Eocene of Europe, but pollen of this genus is known from the middle Eocene to Pliocene of Germany (Krutzsch 1962(Krutzsch , 1970aSontag 1966;Krutzsch & Lenk 1973;Menke 1975;Thiele-Pfeiffer 1980;Gastaldo et al. 1998), the late Eocene of UK (unpublished data, F. Grímsson), the late Oligocene to Miocene of Poland and Austria (Stuchlik 1964;Ziembińska-Tworzydło 1974;Oszast & Stuchlik 1977;Hochuli 1978;Ziembińska-Tworzydło & Ważyńska 1981;Rębas 1985;Stuchlik et al. 1990;; Kohlman-Adamska 1993; Ziembińska-Tworzy-   Stuchlik et al. 2014), and the Miocene of Hungary (Nagy 1969(Nagy , 1985, Turkey (Bouchal 2019 , and the pollen from different extant species are hard to distinguish except in some few taxa. If this was the case in fossil species each morphotype could easily represent more than a single biological taxon. There are not many reports on Arceuthobium macrofossils worldwide. The few Palaeogene records are fragments of shoots and fruiting inflorescences originating from the late Eocene Baltic amber (Sadowski et al. 2017b). The only Neogene macrofossil records are shoots, flowers and fruits, from the late Miocene of Poland (Łańcucka-Środoniowa 1980). Interestingly, the Baltic amber seems to encompass six different co-occurring Arceuthobium species (see Sadowski et al. 2017b). Such a diversity hot spot is only known from current-day Mexico, established following the Miocene (Hawksworth & Wiens 1996). The high number of fossil species in the Baltic amber raises the question whether sexual dimorphism, that is apparent in extant taxa (Hawksworth & Wiens 1996), played a role and lead to an overestimation of the actual palaeo-diversity. Extant Arceuthobium are very host selective and exclusive, raising the additional question whether this was also the case during the Eocene and whether the Baltic amber forests had sufficient host numbers to satisfy at least six different parasites. Sadowski et al. (2017a) note a considerable variety of conifers (at least 12 different taxa) thriving in the Baltic amber forests, including Calocedrus, Quasisequoia, Taxodium (Cupressaceae), Cupressospermum (Geinitziaceae), Abies, Cathaya, Nothotsuga, Pseudolarix, Pinus (Pinaceae), and Sciadopitys (Sciadopityaceae). Finding an appropriate host was apparently not a problem and the high number of different conifers could easily have sustained a diverse Arceuthobium flora. The same can be said about the Miocene Lavanttal flora:  reported at least 20 different conifers, including various Cupressaceae (5 spp.), Pinaceae (14 spp.) and Sciadopitys.
Comparing primitive versus advanced characteristics of extant Arceuthobium, Hawksworth and Wiens (1972) hypothesised that this genus originated in north-eastern Asia during the early Cainozoic. Arceuthobium apparently then dispersed westwards to the Mediterranean area, south into Africa, and north-eastwards across the Bering land bridge into the Americas. Following first molecular evidence, Hawksworth and Wiens (1996) adjusted their theory by adding a possible migration event across the North Atlantic land bridge during the early Cainozoic and constraining the migration across the Bering land bridge as a second event during the Miocene. This would imply that American Arceuthobium originated from two different stocks, an Eocene western Eurasian stock and a Miocene East Asian stock. This contradicts the current molecular phylogenetic framework of the genus by Nickrent et al. (2004) supporting a monophyletic origin of all American taxa with A. azoricum resolved as sister to the American clade, i.e. a single colonisation event via the North Atlantic. The current fossil record suggests a middle Eocene European origin for Arceuthobium (see earlier). Likely, Arceuthobium dispersed southwards into Africa already during the Eocene, similar migration routes have recently been proposed for other plant groups, e.g. the Loranthaceae (from Asia; Grímsson et al. 2017aGrímsson et al. , 2018b and the Picrodendraceae (from Europe; Grímsson et al. 2019). The restricted present-day distribution of Arceuthobium in Africa is most likely the result of post-Miocene climate and vegetation changes. How, if and when Arceuthobium migrated across the North Atlantic land bridge is uncertain. Palynological data place the genus at the gate of the corridor during the late Eocene (UK record, unpublished data, F. Grímsson), but studies on Palaeogene floras of Greenland hold no such records (Grímsson et al. 2014(Grímsson et al. , 2015b2016b, 2016c, 2018a. Also, the Miocene floras of Iceland, both macrofloras and microfloras, have been studied in detail and yield no records of Arceuthobium, but Viscum is present (e.g. Grímsson et al. 2005Grímsson et al. , 2007aGrímsson et al. , 2007bDenk et al. 2005Denk et al. , 2010Denk et al. , 2011Grímsson & Símonarson 2008). The earliest American fossil records of Arceuthobium are from the Miocene (summarised by Hawksworth & Wiens 1996). The fossil pollen from China noted herein represent the only known record of this genus from East Asia. If the hypothesis holds that Arceuthobium originated in Europe during the Eocene, it would be unlikely that it migrated into Asia before the closure of the Turgai Seaway that took place during the early Oligocene (Akhmetiev & Reshetov 1996). It is more likely that Arceuthobium only reached Asia following its closure in the Oligocene and/or Miocene, at the same time as numerous other plant groups dispersed between these continental areas (e.g. Akhmetiev & Reshetov 1996;Manchester 1999;Manchester et al. 2009a;Denk et al. 2012;Grímsson et al. 2012a).
Ecological implications. -Arceuthobium is a small genus with a mostly northern hemispheric distribution. Hawksworth and Wiens (1996) recognised 42 spp., but on the background of molecular phylogenetic data Nickrent et al. (2004) reduced their number to 26. Of these, a single species occurs in Europe, A. oxycedri (DC) Bieb., extending from Spain eastwards through Mediterranean Europe and Africa (Morocco, Algeria) to the Middle East and the Himalayas of India and China (Hawksworth & Wiens 1996;Polhill & Wiens 1998). This species along with A. juniperi-procerae Chiovenda and A. tibetense H.S Kiu et W. Ren are considered the most basal/primitive within the genus (Nickrent et al. 2004). Arceuthobium juniperiprocerae is confined to the highlands of Eritrea, Ethiopia and central Kenya (Polhill & Wiens 1998). Arceuthobium tibetense is only known from the Mainling area in eastern Xizang, Tibet (Hawksworth & Wiens 1996). These three species are placed in Subgenus Arceuthobium Section Arceuthobium and were placed as sister lineages to the clade composing the remaining Asian taxa, A. chinense Lecomte, A. minutissimus JD Hooker, A. pini Hawksws. et Wiens and A. sichuanense (H.S. Kiu) Hawksw. et Wiens of Section Chinense (Nickrent et al 2004). Arceuthobium minutissimum is known from the Himalayas in Pakistan, India, Nepal, and Bhutan, but the other three Asian taxa (A. chinense, A, pini and A. sichuanense) are restricted to south-western China (Hawksworth & Wiens 1996). In addition to these Old World taxa, A. azoricum Hawksw. et Wiens, an endemic to the Azores, occurs on the islands of Terceira, San Jorge, Pico, and Faial in the North Atlantic Ocean (Hawksworth & Wiens 1996). All remaining Arceuthobium (18 spp.) are found in the Americas, occurring in Canada (3 spp.), United States (8 spp.), Mexico (12 spp., greatest diversity), Hispaniola (1 sp.) and continental Central America (3 spp.) (Hawksworth & Wiens 1996;Nickrent et al. 2004).
Arceuthobium are stem hemiparasites that exclusively parasitise conifers and have been found on Abies, Cupressus, Juniperus, Keteleeria, Larix, Picea, Pinus, Pseudotsuga and Tsuga (Hawksworth & Wiens 1972, 1996Polhill & Wiens 1998;Kuijt 2015). Individual taxa can be very selective and parasitising only a single host species, some are found on several different conifers but usually do not cross genera. Dual parasitism (two Arceuthobium species infecting a single host tree) is extremely rare and they rarely parasitise trees that are already occupied by other parasitic plants. It is also believed that climate is a limiting factor in the distribution of Arceuthobium, especially when they do not occur throughout the whole range of their host (Hawksworth & Wiens 1996). At present, Arceuthobium is mostly absent from all equatorial and polar climates (A-and Eclimates) and thrives well in the drier variants (summer or winter-dry) of the warm temperate climates (Cs, Cw), arid climates (B-climates) and snow climates (D-climates). Being a parasite, Arceuthobium is depended on its host. Based on the rich conifer flora previously documented from Lavanttal by , Arceuthobium could have been infecting trees both in the lowland wetlands as well as in the surrounding hills and mountains.  (Gregor 1982). The few additional Eurasian Neogene macrofossil records representing the family are summarised in Mai (1995b). The earliest pollen records are from the Maastrichtian of Canada (Srivastava 1969) and the United States (Nichols 2002). From the Palaeogene, pollen of this family is more 146 F. Grímsson et al. often reported and is known from the Paleocene to Eocene of the United States Bouchal et al. 2016a), the Eocene of Central and East Asia (Wang & Zhao 1980;Hoorn et al. 2012), and the Eocene of western Eurasia (Akkiraz et al. 2006;Worobiec & Gedl 2018). From the Neogene onward Amaranthaceae pollen is very common in Eurasia (summarised by Muller 1981;Stuchlik et al. 2009). Fossil Amaranthaceae pollen is commonly assigned to the form-genus Chenopodipollis Krutzsch (e.g. Stuchlik et al. 2009).
-The Amaranthaceae (including Chenopodiaceae) comprise about 170 genera and over 2400 species. The family has a cosmopolitan distribution, but Amaranthaceae s. str. (excluding the Chenopodiaceae) are mainly tropical and the former Chenopodiaceae are subtropical to temperate. The plants are mostly annual or perennial herbs, subherbs or shrubs, rarely lianas or trees (e.g. Townsend 1993;Kühn 1993). Most Amaranthaceae s. str. occur in regions with low humidity and rainfall, some taxa are aquatic, others occur in inundated or damp depressions, and some are found near water in montane forests (Townsend 1993). The Chenopodiaceae can be dominant components in both marshes and (semi-)deserts. Chenopodiaceae prefer coastal or xeric habitats, they are light demanding plants, and many of them have a dominant position within certain vegetation units. Chenopodium and Atriplex, with non-pronounced xeromorphic attributes, are typical members of vegetation occurring on sandy or stony soil at riverbanks or rocky habitats (Kühn 1993). Based on the cosmopolitan distribution of the family and the overlapping pollen morphology between taxa, potential modern analogues for the fossil Amaranthaceae gen. et spec. indet. 1 and 2 cannot be established. Nonetheless, the fossil pollen grains likely originated from herbaceous plants growing in open and sunny areas, maybe on sandy/stony banks of streams running into the lowland wetlands, or on rocky outcrops in the surrounding highlands. Remarks. -The Amaranthaceae gen. et spec. indet. 2 is much larger than the indet.1 type pollen, the pori are also much wider (1.3-1.5 vs 2.4-2.6), and the density of nanoechini is much higher in the previous taxon than the latter.
Fossil record. -The macrofossil record of Caryophyllaceae is sparse. A fossil inflorescence, Caryophylloflora paleogenica G.J.Jord. et Macphail, with in situ pollen, is known from the Eocene of Tasmania, Australia (Jordan & Macphail 2003). Fossil seeds, assigned to Hantsia, are described from the Eocene of England, UK (Chandler 1960;. All other macrofossils are post-Miocene seed records, assigned to Arenaria, Cerastium, Cucubalus, Minuartia, Silene, and Stellaria (e.g. Zazula et al. 2005;Thompson et al. 2011;Huang et al. 2013). The palynological record of Caryophyllaceae is summarised in Muller (1981), Jordan and Macphail (2003), and Stuchlik et al. (2009 ). They also occur in moist temperate forests or meadows, moist areas in tropical Afromontane regions, but are absent from tropical lowland rain forests (Bittrich 1993). Based on the cosmopolitan distribution of Caryophyllaceae, and the fact that the fossil Lavanttal pollen cannot be assigned to a particular genus (due to overlapping in pollen morphology), further speculations regarding climate preferences of potential modern analogues are impractical. It is likely that the fossil pollen originates from herbaceous plants that were part of open habitats, maybe meadows or shrublands at forest margins, or formed part of the understory in the rich highland forests surrounding the lowland wetlands.
Caryophyllaceae gen. et spec. indet. 2 ( Figure 10G- Remarks. -This pollen type differs from the previously described not only in size of the pollen, but also in the size and shape of echini and the number and conspicuousness of perforations. Also, the echini on the opercula of the Caryophyllaceae gen. et spec. indet. 2 are often fused to elongated, a feature not observed in pollen of the indet. 1 type. This pollen type is morphologically similar (pore membrane configuration) to the 'Silene vulgaris type' of Punt and Hoen (1995).
-Persicaria comprises about 150 species of annual or perennial herbs. It is a cosmopolitan genus, occurring mostly in the Northern Hemisphere with some taxa extending into subtropical and tropical regions, from sea-level to high elevation (Brandbyge 1993;Heywood et al. 2007).
Persicaria plants usually occur near or in the water and are found along the coastline, at riversides, along streams and river banks, along the shoreline of lakes, in swamps, fens, ponds, bogs, tidal marshes, wet ravines, marshlands, moist prairies, on floodplains, and other types of wetlands. The plants can be part of various different forest types (including deciduous forests, mixed deciduous and evergreen forests, and evergreen forests), but are typically concentrated in openings and/or at the forest margins (e.g. Li et al. 2003;Freeman & Reveal 2005;Kantachot et al. 2010;Li 2014;Liang & Li 2014). For a detailed account on the ecology and habitat of P. amphibia consult Partridge (2001). The cosmopolitan distribution of Persicaria and the similarities in pollen morphology of its species, render fossil Persicaria useless for climate inferences using potential modern analogues. The fossil Persicaria pollen from Lavanttal probably originates from herbaceous plants, growing in or close to moving or stagnant water as part of lowland wetlands or in open areas of the forests surrounding the basin, at forest margins and/or along streams.
-Rumex comprises about 195 species of perennial or annual herbs (POWO 2017). Like the other found Polygonaceae, it is a cosmopolitan genus but with a preference of the temperate regions of the world (Brandbyge 1993). Rumex occurs from sea-level to an elevation of about 4300 m. The plants are found in various environments (e.g. coastal, alluvial and montane habitats), including, swamps, marshes, bogs, shores of lakes, banks of streams and rivers, grasslands, (wet) meadows, (moist) valleys, (dry) mountain slopes, forest margins, sandy and gravelly shores, sandy planes, sand dunes, rocky outcrops, rocky fissures, and even saline deserts and sands (Li et al. 2003;Freeman & Reveal 2005). As in the case of other Polygonaceae, it is useless for climate reconstruction. Based on the variable habitats of extant Rumex it is uncertain if the pollen originates from plants that were growing in the lowland wetlands or suitable patches in the highland and/or mountain forests surrounding the basin. Remarks. -The pollen morphology of Alangium has been studied quite thoroughly by Reitsma (1970), but see also Chao (1954), Straka et al. (1967), Eyde et al. (1969), Eyde (1972), Cerceau-Larrival et al. (1984) and Martin et al. (1996). These palynological studies were based on the c. 20 taxa known at that time. Recent revision on three of the four currently accepted sections within Alangium (sections Alangium, Conostigma, Rhytidandra) by de Wilde and Duyfjes (2016 shows that they comprise 43 species. Section Marlea has not been revised so far but is believed to hold less than 10 taxa (de Wilde & Duyfjes 2016. The pollen morphology of Alangium is therefore in need of revision to clarify the characteristics of each section and to assess whether important and/or diagnostic features are overlapping between sections. The fossil Alangium pollen from Lavanttal is heterobrochate-reticulate to striatoreticulate and with narrow muri. Until now, Alangium kurzii Craib (section Marlea) is the only extant taxon known to produce comparable pollen (Reitsma 1970, plates 7-10).
Ecological implications. -Alangium kurzii is a deciduous shrub or tree that can be up to 28 m tall. It occurs from 50-1600 m a.s.l. and has a vast distribution across southeast Asia (including Myanmar, Thailand, Vietnam, Malaysia, Indonesia and the Philippines) and East Asia (including southern and eastern China, Korea and Japan) (Bloembergen 1935(Bloembergen , 1939Qin & Phengklai 2007). In China, Alangium kurzii is a woodland tree occurring at 600-1600 m a.s.l. in mixed 152 F. Grímsson et al.
deciduous-evergreen broadleaved and conifer forests (Qin & Phengklai 2007) in fully humid to winter-dry temperate climates with hot summers (Cfa, Cwa; meridio-nemoral vegetation element; File S1) with mean annual temperatures ranging from 11.7 to 24.8°C, an annual precipitation of 711 to 2435 mm, and coldest month mean temperatures of 1.1 to 20.1°C (Fang et al. 2009). Based on the preferred habitat of extant Alangium kurzii, and the rareness of this fossil pollen type, it is likely that the Lavanttal Alangium was not growing in the lowland wetlands, but was part of the forest community surrounding the basin, and probably growing along streams or in sparse forest units on well-drained substrates.
Genus Cornus L. Cornus sp. Remarks. -The pollen morphology and ultrastructure of Cornus has been studied by Ferguson (1977). For additional LM-and SEM-based work on Cornus pollen see e.g. Adams and Morton (1976), Stafford and Heath (1991), Chester and Raine (2001), Perveen and Qaiser (2002), Mert (2009), Li et al. (2011), Miyoshi et al. (2011), and Kilie and Tuttu (2017. In his study on c. 44 different Cornus species, Ferguson (1977) concluded that all taxa produce similar pollen (stenopalynous genus sensu Halbritter et al. 2018) but divided them into two groups: (a) 'Cornus mas-subtype', and (b) 'Cornus sanguinea-subtype'. The main difference between these two subtypes is the polar/equatorial (P/E) ratio. Pollen assigned to the Cornus mas-subtype are ± isodiametric, and those grouped in the C. sanguinea-subtype are ± prolate.  (Manchester et al. 2009b). The complete post-Paleocene fossil record of Cornus was summarised in detail by Eyde (1988), including the alleged pollen record. Both Eyde (1988) and Muller (1981) were sceptic about all the fossil Cornus pollen that had been published/ described (using LM only) at that time, stating they lack diagnostic features to differentiate them from pollen grains of other Cornaceae and related groups. The earliest fossil Cornus pollen studied using SEM is reported from the late Paleocene of Almont, North Dakota, USA .   These include mixed broad-leaved and conifer forests, mixed thickets/woods, dense forests, the margin of woods, and scrub vegetation. Cornus is frequently growing along streams in both lowlands and at higher elevations, on hillsides and in real mountainous regions (Xiang & Boufford 2005). In China, most Cornus species occur in fully humid warm temperate climates with hot summers or winter-dry warm temperate climates with hot to warm summers (Cfa, Cwa, Cwb; meridio-nemoral vegetation element; File S1). Some Cornus extent into winter-dry snow climates with hot to warm summers or cool summers and cold winters (Dwa, Dwb, Dwc; nemoral vegetation element; File S1). In North America, Cornus grow from sea-level to c. 3400 m a.s.l. Cornus plants are found in both open vegetation as well as in dense forest, in mesic or dry-mesic deciduous hardwood forests, alluvial woods, or dry woodlands. As in China, Cornus in America often occurs along river and stream banks, as well as near wet meadows, swamp margins and marshes (Murell & Poindexter 2016). North American Cornus species occur in fully humid warm temperate or snow climates with hot or warm summers (Cfa, Cfb, Dfa, Dfb; nemoral vegetation element; File S1). Based on present-day Cornus, the Lavanttal Cornus pollen represents a nemoral or meridio-nemoral vegetation element originating from plants growing in the lowlands, at the boundary of swamps or on levees, or from plants growing along streams in highland areas surrounding the basin.
Remarks. -Pollen of Nyssa has been studied in detail by various authors using both LM (Sohma 1963(Sohma , 1967 and SEM (Saito et al. 1992;Göschl 2008;Li et al. 2011). Nyssa aquatica L., N. ogeche W.Bartram ex Marshall, N. sylvatica Marshall and N. sinensis Oliv. produce pollen with less sculptured or psilate areas surrounding the colpi. In N. javanica (Blume) Wangerin and N. bifida Craib a distinct nanorugulate sculpture is present in areas adjacent to the colpi (Göschl 2008). Information concerning this character state in N. talamancana Hammel et N. Zamora is currently unavailable. All investigated Nyssa pollen from Lavanttal displayed no nanorugulation in the colpus area.
Fossil record. -The fossil record of Nyssa has been summarised by Eyde (1997) and Manchester et al. (2015). Early macrofossil records of Nyssoideae (fruits) date back to the Upper Cretaceous (early Coniacian) of Japan (Takahashi et al. 2002). The three sculpture types known from endocarps of extant species (ridged with sunken bundles, ridge raised bundles, and smooth) have been reported from the early Eocene of North America and Europe, indicating radiation prior to the Eocene. The form-genera Nyssapollenites Thiergart ex R.Potonié and Nyssoidites R.Potonié, Thomson et Thiergart ex R.Potonié have commonly been used for fossil Nyssa-like pollen. The Cainozoic palynological record of Nyssa is summarised by Muller (1981) and Eyde (1991), and most recently by Stuchlik et al. (2014), accepting earliest records from the Paleocene of the Northern Hemisphere.
Ecological implications. -The nine extant species of Nyssa show a disjunct distribution, with five species occurring in eastern North America (N. aquatica, N. biflora Walter, N. ogeche, N. sylvatica, N. ursina Small), one in Central America (N. talamancana), and three in east to southeast Asia (N. bifida, N. javanica, N. sinensis) (Wen & Stuessy 1993;Tucker 2016;POWO 2017). All extant species of Nyssa are deciduous trees. The North American species are commonly found in habitats with water-logged soils (swamps, floodplain forests, riparian forests) at low elevations in fully humid warm temperate climate (Cfa). Nyssa sylvatica shows the widest ecological range, thriving in wet, well-drained, and even dry environments/habitats, from subtropical to snow climates (Aw, Cfa, Cfb, Dfa, Dfb; meridio-nemoral vegetation element; File S1), at an elevation from sea level up to 1100 (1600) m, and frost resistance in mature trees to −30°C (Sakai & Weiser 1973;Tucker 2016). Nyssa talamancana thrives at middle elevations in Costa Rica and Panama under fully humid equatorial climate or fully humid warm temperate climate with warm summers (Af, Cfb; tropical-oreotropical vegetation element; File S1) together with other relict genera, e.g. Ticodendron, Alfaroa, Oremunnea and Gordonia (Hammel & Zamora 1990). Nyssa sinensis is found in wet mixed forests along streams at elevations from 300 to 2700 m (Haining & Chamlong 2007), under fully humid to winter-dry warm temperate climate with hot to warm summers (Cfa, Cfb, Cwa, Cwb; meridio-nemoral vegetation element: File S1). The habitats of extant Nyssa species producing pollen similar to the fossil from Lavanttal point to a nemoral to meridio-nemoral vegetation element. Its pollen probably originated from plants growing in the lowlands, in swamps, on waterlogged soils, or as riparian elements.

Order Ericales Bercht. et J.Presl
Family Ebenaceae Gürke Genus Diospyros L. Diospyros sp. Pollen of extant Diospyros species display a wide range in size and outline, but all have a (micro) rugulate sculpture (Geeraerts et al. 2009). The relatively long colpi and perforations observed in interapertural areas (SEM) of the fossil pollen is shared with pollen of many extant Diospyros taxa (Geeraerts et al. 2009, tables I and II).
Fossil record. -The macrofossil record of Diospyros has been summarised by Hiern (1873), Berry (1912), and Basinger and Christophel (1985). Both Basinger and Christophel (1985) and Wallnöfer (2001Wallnöfer ( , 2004 (Wallnöfer 2001(Wallnöfer , 2004. Diospyros are mostly small to medium-sized evergreen, less frequently deciduous, trees in the forest understorey. The trees often thrive along rivers but can also be found in swamps and periodically flooded environments. Some species are part of well-drained forests and occur in deciduous forests (e.g. D. lotus L., D. kaki L.f., D. virginiana L.); some are even growing in fire-prone savannahs (Wallnöfer 2001(Wallnöfer , 2004. In North America, D. virginiana, is a widely distributed tree (up to 40 m tall) in the eastern United States. It occurs from 0 to 1100 m a.s.l. and is found in various forest types, from seasonally flooded bottomlands to dry ridgetops (Eckenwalder 2009), predominantly thriving in fully humid warm temperate 156 F. Grímsson et al. climates with hot to warm summers (Cfa; meridionemoral vegetation element; File S1) and extending into Dfa climates with coldest month minima of −4.9° (Thompson et al. 1999). The other North American taxon, D. texana Scheele, distributed in Texas and Mexico, occurs from 0 to 1800 m a.s.l., and is part of open lowland bottomlands, prairie margins, and is found on rocky hillsides. In Texas, it thrives under fully humid warm temperate climate with hot summers (Cfa), but in the southern part of its distribution area, Mexico, it is found in hot arid steppe climate (BSh; File S1). China, mostly its south-eastern and south-western parts, is home to about 60 species of Diospyros (Lee et al. 1996). Small shrubs to large trees (up to c. 25 m tall), which can be either deciduous or evergreen, which occur from sea level to an elevation of 2700 m. The plants are part of both deciduous and broad-leaved evergreen forests, as well as mixed broad-leaved evergreen-deciduous forests. They often occur in forested ravines or on slopes, and in forests beside streams or moist lowland valley forests (Lee et al. 1996). The Chinese Diospyros fall into two major groups regarding climate preferences. About onethird of the taxa grow under fully humid to winterdry warm temperate climate with hot to warm summers (Cfa, Cfb, Cwa, Cwb; nemoral to meridionemoral vegetation element; File S1). About twothirds of the species either extend into (tropicalmeridional vegetation element) or are confined to various equatorial climates (tropical vegetation element), like fully humid rainforest (Af), monsoon (Am), or savannah climates with dry winter (Aw; File S1). Based on the extant habitats of warm temperate Diospyros species, the fossil pollen from Lavanttal most likely represent shrubs or small trees, characteristic for the understory. The pollen could originate from plants growing in the lowland wetlands or along streams at the periphery of the basin. It could also derive from plants growing further away from the accumulation zone, representing ravine vegetation or a highland element.
( Figure 13A- Fossil record. -It is not easy to affiliate fossil Ericaceae tetrads to extant lineages and/or genera. Most fossil records are documented using LM only and assigned to different species of the form-genus Ericipites Wodehouse (e.g. Stuchlik et al. 2014). We are unaware of any convincing pre-Quaternary Andromeda macrofossil or pollen record.
Ecological implications. -Andromeda polifolia is the single modern species of genus Andromeda (POWO 2017). It is a low growing evergreen shrub that is usually 10 to 25 cm tall but can grow up to 40 cm. This species is present in all circumboreal regions of the Northern Hemisphere. It inhabits wet sites, (peat) bogs, fens, swamps, margins of pools, and boggy shores, from sea level up to 1500 m, throughout boreal forests and the Arctic (Jacquemart 1998;Fabijan 2009). For a detailed summary of the distribution and the ecology of A. polifolia see Jacquemart (1998). Andromeda polifolia does not extend into tropical (A) or arid (B) climates but is found in fully humid warm temperate and snow climates with hot to cold summers (Cfa, Cfb, Cfc, Dfb, Dfc; File S1), summer-and winter-dry snow climates (Dwb, Dwc, Dsb, Dsc) and Tundra climates (ET) (boreal to nemoral vegetation element; File S1). During the Miocene of Lavanttal Andromeda probably was part of the lignite forming swamp community or growing at the margins of conifer forests belts at higher elevation.
Remarks. -Arbutus has large tetrads compared to most Ericaceae. The colpi are distinct, but the boundaries between individual pollen grains (in mesocolpium area) are extremely vague (linear perforations) and untypical for this family. Individual pollen grains are ± triangular in polar view (LM) versus lobate to circular in most Ericaceae. This, in combination with the granulate and perforate sculpture observed with SEM, clearly place these fossil tetrads from Lavanttal in Arbutus. The pollen morphology of most Arbutus species have been documented in detail using both LM and SEM by e.g. Adams and Morton (1979), Lewis et al. (1983), Warner and Chinappa (1986), Barento et al. (1987), Luteyn et al. (1995), Sarwar (2007), Sarwar et al. (2008), andHalbritter (2016a).
Fossil record. -The fossil record of Arbutus is meagre. In the United States, fossil Arbutus leaves have been reported from the early and late Oligocene of Colorado (Axelrod 1987;Gregory & McIntosh 1996), the Miocene of Nevada and Oregon (e.g. Axelrod 1991;Graham 1999), and the Miocene/ Pliocene of California (Axelrod 1950). In Europe, fossil Arbutus leaves are known from the Eocene to Pliocene (summarised in Mai 1995b). Noteworthy is that genetic investigations of Arbutoideae have shown that European Arbutus species are sister to all American genera (including Arbutus) of this subfamily (Hileman et al. 2001). As mentioned previously, it is not easy to affiliate fossil Ericaceae pollen tetrads to extant lineages and/or genera, therefore, most fossil Ericaceae pollen tetrads are assigned to the form genus Ericipites (e.g. Stuchlik et al. 2014). There seems to be no previous convincing pre-Quaternary Arbutus pollen record.
Ecological implications. -Arbutus is a small genus, comprising c. 14 species of shrubs and trees, occurring in western Europe to the Mediterranean, Macaronesia, and western Canada to Central America (Hileman et al. 2001;Sørensen 2009;POWO 2017 (Hileman et al. 2001;Sørensen 2009;POWO 2017). Arbutus arizonica thrives in riverine forest and along seasonally moist waterways, at an elevation of 1500 to 2400 m, in Arizona and northern Mexico. Arbutus menziesii occurs in open forests, on rocky slopes, foothills, in ravines, and along shores, at an elevation of 0 to 1800 m, from British Columbia to northwest Mexico. In tropical Central America Arbutus is found primarily in forested montane areas dominated by Pinus and Quercus (Sørensen 2009). The American Arbutus species grow under a wide range of climates, from fully humid temperate conditions with warm to cool summers (Cfb, Cfc) to summer-or winter-dry climates (Csb, Cwa, Cwb), and even extend into equatorial and arid climates (Aw, BSh, BSk). The Arbutus pollen from the Miocene of Lavanttal could have originated from trees growing along streams or in ravines at the periphery of the basin, or from drier hill sites or mountain areas surrounding the lowlands. Remarks. -The pollen morphology of Empetrum has been studied with both LM and SEM and figured, among others, by Warner and Chinappa (1986), Foss and Doyle (1988), Beug (2004), Sarwar (2007), Miyoshi et al. (2011), andHalbritter (2016b). The fossil pollen from Lavanttal corresponds in size and morphology to extant Empetrum pollen, and the sculpture observed in SEM is comparable to that of E. nigrum L. as depicted by Halbritter (2016b).

Genus
Fossil record. -The few North American and European macrofossil records of Empetrum are mostly confined to the Quaternary (e.g. Mai 1995b;Graham 1999). Still, a single endocarp assigned to this genus is reported from the middle Miocene of Denmark (Friis 1979). According to Mai (1995b), Empetrum pollen is first recovered from the Miocene of Europe, but then became a prominent component in Pliocene and younger peat deposits of that region. As noted earlier, Ericaceae pollen tetrads are very similar in morphol-ogy and it is not easy to affiliate fossil tetrads to extant taxa unless observed with SEM. Most fossil Ericaceae tetrads studied using LM are assigned to the formgenus Ericipites (e.g. Stuchlik et al. 2014). There are currently no previous pre-Miocene Empetrum pollen records verified using SEM.
-Empetrum composes about four extant species of prostrate shrubs (Murray et al. 2009;POWO 2017). Three of the species are restricted in distribution. Empetrum rubrum Vahl ex Willd. is native to southern South America and the Falkland Islands, where it thrives under fully humid warm temperate climates, with warm to cool summers and cold winters, and extends into summer-dry as well as tundra climates (Csb, Cfb, Cfc, ET; File S1). Empetrum atropurpureum Fernald et Wiegand and E. eamesii Fernald et Wiegand are native to eastern Canada and north-eastern United States, where they occur at an elevation from sea level to c. 1500 m, on dunes, sandy terraces, coastal rock barrens, alpine heath, and exposed mountain slopes near treeline (Murray et al. 2009). Both species grow under fully humid snow climates with warm to cool summers (Dfa, Dfb, Dfc; File S1) with E. atropurpureum extending into fully humid temperate climates (Cfb) touching Cfa at the coast of Maine, USA. The most common species, E. nigrum, has a circumpolar (temperate to Arctic) northern hemispheric distribution (POWO 2017). According to Murray et al. (2009), E. nigrum occurs in North America, Greenland and Europe at elevations from sea level to c. 1900 m, in windswept southern arctic and alpine tundra and open subalpine and boreal forests and mountain summits, as well as on exposed, coastal bluffs and in sphagnum bogs. In China (East Asia), E. nigrum can be found in forests, on stony hills, at elevations of 700 to 1500 m (Fang et al. 2005). Empetrum nigrum does not extend into tropical (A) or arid (B) climates but is found in fully humid warm temperate and snow climates with hot to cold summers (Cfa, Cfb, Cfc, Dfb, Dfc; File S1), summer-dry warm temperate climate with warm summers (Csb; File S1), summer-to winter-dry snow climates (Dsc, Dwb, Dwc), and Tundra climates (ET) (boreal to nemoral vegetation element; File S1). It is possible that during the Miocene of Lavanttal, Empetrum was growing on shady slopes or in shadows at the edge of woodlands in the hillsides and/or mountains surrounding the Lavanttal Basin.
Remarks. -Pollen in permanent tetrads showing verrucate to microverrucate sculpture and nanoechinate suprasculpture (SEM) are typical for Erica. Despite the number of publications including Erica pollen, only a small portion, less than 100 species, are palynologically studied so far (e.g. Oldfield 1959;Foss & Doyle 1988;Díez & Fernandes 1989;Mateus 1989;Oliver 2000;Sarwar 2007;Sarwar & Takahashi 2014;Wrońska-Pilarek et al. 2018; and references cited therein). Based on the taxa studied so far, Erica pollen can be divided into two major groups: (1) pollen dispersing in permanent tetrahedral tetrads (majority), and (2) pollen dispersing as monads (minority). All the Erica type pollen from Lavanttal belong to the former. Because of the numerous unstudied extant taxa and the overlapping pollen morphology of the studied ones, we refrain from affiliating the Lavanttal pollen tetrads to any particular species group or intrageneric lineage.
Fossil record. -According to Mai (1995b, p. 182), the few pre-Quaternary leaf records of Ericoideae/ Erica are doubtful and not based on reliable leaf anatomy nor cuticle analyses. Rare fruits/capsules of Erica are documented from the late Miocene of Europe (van der Burgh 1987). As with other fossil Ericaceae pollen tetrads, they are hard to affiliate to extant lineages and/or genera unless studied with SEM. Therefore, most fossil Erica tetrads are lumped into the form-genus Ericipites (e.g. Stuchlik et al. 2014). The earliest pollen records of Erica, verified using combined LM and SEM, are from the early Miocene of Europe, but they become common elements in late Miocene and younger sediments of that region (e.g. Zetter 1991;Ferguson et al. 1998;van der Burg & Zetter 1998;Hofmann et al. 2002).
Ecological implications. -Erica is one of the largest plant genera, comprising c. 858 species (POWO 2017) of perennial woody prostrate shrubs and trees, up to 10 m tall (Oliver 2000;Stevens et al. 2004). Erica species show a narrow north-south geographic distribution, spanning north-western Europe and the Mediterranean region (c. 23 spp.), the Middle East (the genus extends eastwards into Turkey and Lebanon and the south-western part of the Arabian Peninsula), eastern and soutern Africa  species that is widespread in both Europe and Africa. Erica arborea is present in fully humid to summer-dry temperate (Cfa, Cfb, Csa, Csb; File S1) climates in Europe. In Africa, this species is found in winter-dry to fully humid temperate climates in mountainous regions (Cfb, Cwa, Cwb; File S1). In Europe, Erica are part of different shrub communities, and in Africa they are part of montane scrub or grasslands (Quezel 1978;Oliver 2000;McGuire & Kron 2005). In general, Erica thrives under winter-dry, summer-dry, and fully humid temperate climates with hot or warm summers (Cfa, Cfb, Csa, Csb, Cwa, Cwb; File S1), as well as fully humid snow climates with warm to cool summers (Dfb, Dfc), but rarely extends into equatorial (A) or arid (B) climates. The fossil Erica pollen tetrads from Lavanttal either originate from small shrubs that occurred in open habitats (dry to wet substrate, lowland to highland) or as understory growth (ground vegetation) in woodlands.
Remarks. -The Erica sp. 2 pollen tetrads are easily distinguished from the Erica sp. 1 type. In Erica sp. 2 the verrucae around the colpi are fused and forming 'large' rugulae, a feature not observed in Erica sp. 1. Also, the nanoechini in Erica sp. 2 are much smaller and more numerous than in Erica sp. 1. A similar ornamented Erica pollen tetrad is reported from the middle Miocene of Turkey (Bouchal et al. 2016b, figure 20G-I).
Remarks. -The Erica sp. 3 pollen tetrads are similar to those of Erica sp. 2, but are slightly larger, the rugulae are more homogeneous in size and relief, and the nanoechini are also much shorter and blunt.
Remarks. -The Erica sp. 4 pollen tetrads are the only fossil Erica tetrads with clearly uniform nanoverrucate sculpture in SEM.
Remarks. -Pollen tetrads assigned to Erica sp. 5 differ from the remaining Erica sp. 1-4 in the sculpture observed with SEM. The tetrads are nanoverrucate in Erica sp. 5, but verrucate to microverrucate or rugulate in Erica sp. 1 and sp. 2. The sculpture elements around the apertures and is areas of mesocolpium in Erica sp. 5 are of similar size and outline but vary substantially in Erica sp. 2. The Erica sp. 5 pollen tetrads are most similar to Erica sp. 4 but the nanoverrucae are larger in Erica sp. 5, taller, and more globular in outline, and with a higher number of nanoechini per nanoverruca.   Remarks. -Pollen tetrads of this and the following type definitely belong to the Ericaceae, but we are currently unable to affiliate them with certainty to a particular genus. They could also represent extinct lineages (e.g Kowalski & Fagúndez 2017).
Ecological implications. -Pouteria comprises c. 320 species of trees and shrubs occurring in the Americas (c. 200 spp.), in Africa (c. 5 spp.), and in con-tinental Asia, Malesia, Australia and the Pacific (c. 120 spp.) (Pennington 1991(Pennington , 2004. Pouteria is a pantropical genus occurring in wet lowland (rain) forests, lowland swampy forest, (semi-)evergreen lowland forests, periodically flooded forests, upland (rain)forests, (evergreen)montane (rain)forests, riverside gallery forests, along savannah edges, and in dry thickets on limestone hills (Pennington 1990). Since Pouteria are pollinated by insects, the fossil pollen grains likely originated from plants growing close to the depositional site. We hence consider that the Pouteria pollen from Lavanttal originated from sheltered understorey shrubs (or small trees) that were part of the lowland wetland forests.   Remarks. -The fossil Sideroxylon pollen falls within the range of 'Pollen Type 6ʹ (Harley 1991a, figure 11) and 'Subtype 6A to C' (Harley 1991b, figure 24) (Pennington 1991(Pennington , 2004. Sideroxylon is a tropical to subtropical genus, and depending on the species, occurring among others in coastal vegetation, mangroves, wet lowland forests, (lowland) tropical rainforests, montane rain forests, cloud forests, moist seasonal evergreen forests, humid forests, lower montane forests, mixed oak-semievergreen forests, tropical (dwarf) deciduous forests, seasonal (semi)deciduous forests, dry forests, and arid thorn forests (Pennington 1990). Noteworthy is, recent genetic studies indicate that the North American clade of Sideroxylon, including the most frost hardy taxa, already split during the upper Eocene (Stride et al. 2014). Extant Sideroxylon are entomophilous (insect-pollinated) and, therefore, the fossil pollen grains from Lavanttal probably originate from plants growing close to the depositional site, from shrubs or small trees occurring in the lowland wetland forest or along streams at moderate elevation.
Fossil record. -The Sapotaceae pollen record has been summarised by Muller (1981), Harley (1991aHarley ( , 1991b, Song et al. (2004), and Stuchlik et al. (2014). Few of the records date back to the late Cretaceous, but most are confined to the Cainozoic, suggesting a cosmopolitan distribution during the late Eocene. These fossil Sapotaceae pollen grains have been assigned to various species divided into two main form-genera, Sapotaceoidaepollenites R.Potonié, Thomson et Thiergart ex R. Potonié, and Tetracolporopollenites Pflug et Thomson. Most of these records are based on LM observations only but see Harley et al. (1991), and any affiliation to extant lineages and/or genera are uncertain (cf. Harley 1991b). Relying on the monumental work by Harley (1986aHarley ( , 1986bHarley ( , 1990Harley ( , 1991aHarley ( , 1991bHarley ( , 2004   huca comprises c. 100 species that occur from India through Malesia and south China and New Guinea (Pennington 1991(Pennington , 2004. Manilkara is a pantropical genus composing c. 65 species occurring in the Americas (c. 30 spp.), Africa and Madagascar (c. 20 spp.), and in Asia and across the Pacific (c. 15 spp.) (Pennington 1991(Pennington , 2004.
Mimusops comprises c. 41 species that are distributed in Africa (c. 20 spp.), Madagascar (c. 15 spp.), the Mascarene Islands (4 spp.), the Seychelles, and in Asia and the Pacific (one species). The c. 110 spp. of Palaquium range from India through southeast Asia to the Pacific Islands. Xantolis is a small genus comprising c. 14 species ranging from southern India to Vietnam and southern China, with one species in the Philippines (Pennington 1991(Pennington , 2004. All these genera are insectpollinated and occur in tropical or/to subtropical regions, where they are part of various vegetation units. We speculate that the Sapotaceae gen. et spec. indet. 1 and 2. originate from shrubs or small trees that were part of the lowland wetlands or growing along streams in the surrounding hillside forests of the Lavanttal Basin. Sapotaceae gen. et spec. indet. 2 ( Figure 18G-L, 20A-C) Description. -Pollen monad, prolate, outline quadrangular to pentangular in polar view, elliptic in equatorial view; polar axis 37-40 µm long in LM, 34-36 µm long in SEM equatorial diameter 27-32 µm wide in LM, 22-26 µm wide in SEM; stephano(4-5)colporate, colpi narrow, endopori lalongate elliptic, margins of endopori thickened; exine 1.9-2.5 µm thick, nexine thinner than sexine, sexine thickened along colpi (LM); tectate; sculpture scabrate in LM, slightly rugulate, microverrucate, perforate, fossulate in SEM, colpus membrane microverrucate (SEM).
Remarks. -The Sapotaceae gen. et spec. indet. 2 pollen differs from the sp. 1 pollen type in both size and outline in polar view (square versus circular or lobed). The sp. 2 pollen type is also consistently tetra-aperturate, but the sp. 1 pollen is usually equipped with five apertures. Remarks. -Pollen from three out of five Rehderodendron species has been studied using LM and SEM by Liang and Yu (1985) and Morton and Dickison (1992). The sculpture range, in the area of the mesocolpium, of the Lavanttal pollen is comparable to that documented by Liang and Yu (1985) for both R. kweichowense Hu and R. macrocarpum Hu (Liang & Yu 1985, plate 3, figures 22, 23).
Fossil record. -The scarce macrofossil record of Rehderodendron is summarised by Mai (1970) and Manchester et al. (2009a). Fruits of this genus are documented from the early Eocene of UK (Mai 1970), the Miocene of Germany, Poland and Czech Republic (Mai 1970), the Pliocene of France (Geisser & Gregor 1981), Italy (Martinetto 1998) and Romania (Mai & Petrescu 1983). The fossil leaf records of Styracaceae are considered unreliable by Fritsch (2004) because they lack detailed anatomical features such as stellate or scale-like trichomes. Fossil pollen of Rehderodendron, identified using combined LM and SEM, has been reported from the Miocene of Germany (Ferguson et al. 1998) andAustria (Kovar-Eder et al. 1998).
Ecological implications. -Rehderodendron is a small genus comprising five species of deciduous trees (up to 15 m tall) distributed in south-western China, Myanmar and Vietnam (Hwang & Grimes 1996;Fritsch 2004). The plants occur in dense forests, mixed broad-leaved evergreen and deciduous forests, at elevations from 100-1500 m (Hwang & Grimes 1996). All extant Rehderodendron species are currently growing under fully humid to winterdry warm temperate climates with hot to warm summers (Cfa, Cfb, Cwa, Cwb; nemoral to meri-

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F. Grímsson et al. dio-nemoral vegetation element; File S1). Based on the current habitat of Rehderodendron the Lavanttal pollen may represent medium-sized trees that were part of the well-drained dense forests surrounding the basin.
Fossil record. -The fossil record of Symplocos was summarised among others by Krutzsch (1989), Mai and Martinetto (2006) and Tiffney et al. (2018). All authors agree that the leaf record is hard to judge because of their similarity to related genera. The fruit/seed fossil record is fairly rich. It includes finds from the middle Eocene of Oregon (Manchester 1994a) and the early Miocene of Vermont, USA (Tiffney et al. 2018), the Eocene to Pliocene of Germany (Mai & Martinetto 2006;Manchester & Fritsch 2014), the Oligocene to Miocene of Poland, Switzerland and Austria (e.g. Mai & Martinetto 2006), the Miocene of Czech Republic, France and Denmark, (e.g. Mai & Martinetto 2006;Manchester & Fritsch 2014), the Pliocene of the Netherlands, Italy (e.g. Mai & Martinetto 2006) and Japan (Miki 1937 Muller (1981), Krutzsch (1989), Ivanov (1995Ivanov ( , 2004 and Stuchlik et al. (2014). These include doubtful pollen from the Cretaceous of the Americas. The pollen record suggests that Symplocos already had a European-North American distribution during in Eocene. Based on the compiled fossil record, Manchester and Fritsch (2014) and Fritzsch et al. (2015) hypothesised an early Cainozoic European origin of the genus followed by an Eocene dispersal across the North Atlantic land bridge into the Americas. Dispersal from Europe into Asia is believed to have taken place following the closure of the Turgai Strait. There are only a few fossil Symplocos pollen grains that have been studied using combined LM and SEM. These include two pollen types from the Eocene of Profen, Germany (Haring 2014), three pollen types from the late Oligocene/early Miocene of Altmittweida, Germany (Kmenta 2011;Kmenta & Zetter 2013), five pollen types from the Miocene of Kreuzau, Germany (Ferguson et al. 1998, figured only one of the types), and numerous pollen types from the Miocene of Austria (Kovar-Eder et al. 1998, figured one out of three pollen types; Zetter 1998, figured none out of three pollen types; Meller et al. 1999, figured one out of five pollen types). Ashraf and Moosbrugger (1996) figured three types of Symplocoipollenites using SEM from the Lower Rhine Embament of Germany. Therefore, micrographs documenting the SEM-based sculpture variation in fossil Symplocos is lacking. Fossil pollen most similar 172 F. Grímsson et al. to the Lavanttal fossils can be found in the Miocene (Ferguson et al. 1998, plate 6, figures 8-11), and, Eocene of Germany (Haring 2014, plates 8, 9). Lodhra, growing in east to southeast Asia, are evergreen shrubs to small understory trees (mostly less than 10 m tall; rarely up to 30 m tall) occurring in mixed forests, especially on forested slopes, and depending on their geographic occurrence are found at elevations between 100 and 3000 m (Wu & Nooteboom 1996). They grow under equatorial monsoon and winter-dry savannah climates, as well as in fully humid to winter-dry warm temperate climates with hot or warm summers (Am, Aw, Cfa, Cfb, Cwa, Cwb; tropical-meridional to meridio-nemoral vegetation element; File S1). The Mexican to Central American Symplocos of subg. Symplocos sections Symplocos and Hopea (S. culminicola Standl. et Steyerm. and S. longipes Lundell) are also evergreen shrubs to small or medium-sized trees that are usually less than 20 m tall, and rarely up to 50 m in S. hartwegii A.DC. (Kelly et al. 2016). The plants occur mostly in tropical rainforests, from sea level up to 1600 m elevation, and in cloud forests, at elevations between 600 and 3350 m. They are also found in cool mountain forests, mountain rain forests, broad-leaved evergreen forests and mixed forests (Kelly et al. 2016). In this part of the world, Symplocos thrives under fully humid equatorial rainforest climate, equatorial monsoon climate, and summer-dry equatorial savannah climates (Af, Am, Aw; tropical vegetation element; File S1), but extends into fully humid or winter-dry warm temperate cli-mates with warm summer (Cfb, Cwb). Based on the above, the various Symplocos pollen (sp. 1 to sp. 4) could represent evergreen shrubs or small trees that were part of the understory in the mixed deciduousevergreen broad-leaved and conifer forests surrounding the basin.
Symplocos sp. 2 ( Figure 21J- Nagamasu (1989a) and Lieux (1982). concluded that Symplocos originated in Eurasia and dispersed into the Americas during the early Cainozoic. They also postulated that dispersal from North America back to Eurasia occurred within Symplocos sect. Hopea in the middle to late Miocene, pinpointing S. tinctoria as one of two species involved in the Miocene disjunction. This fossil pollen type from the late middle Miocene of Lavanttal seems to collaborate that theory.
Ecological implications. -Symplocos tinctoria is distributed in south-eastern United States, from sea level to an elevation of c. 1400 m. It is a deciduous shrub to small tree and part of several different vegetation units ranging from lowland wetlands to well-drained or dry upland forests (moist mixeddeciduous hardwoods to dry pine-oak woods). It occurs in maritime forests, swamps, hammocks, bottomlands, flatwoods, streamheads, baygalls, on rocky summits and in ravines ). Symplocos tinctoria is growing under fully humid warm temperate climate with hot to warm summers (Cfa, Cfb; meridio-nemoral vegetation element; File S1). The fossil pollen could have originated from small understory trees growing either in the lowland wetlands or in forests on dryer substrates surrounding the Lavanttal Basin.

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F. Grímsson et al. Table I. Angiosperm pollen described in this study in comparison to Klaus (1984).
Based on the dominant climatic preferences of potential modern analogues of the fossil Myrtales to Ericales from the Sarmatian of Lavanttal, as expressed by their 'Köppen signatures' (Figure 23, Table III), equatorial (A [lowlands]), arid (B), and polar (E [at highest altitudes]) climates seem unlikely for this region at that time. Even though some of the taxa, including Reevesia, Pistacia, Zanthoxylum, Nyssa, Dyospyros, Arbutus, Erica, and Symplocus, extend into equatorial climates, they also thrive in various warm temperate (C) climates, while by far the most elements are extra-tropical. The majority of the woody plants, Craigia, Reevesia, Tilia, Pistacia, Zanthoxylum, Acer, Alangium, Cornus, Nyssa, Dios- pyros, Arbutus, Erica, Rehderodendron, and Symplocos, are mostly nemoral and/or meridio-nemoral vegetation elements and typical of warm temperate (C) climates ( Figure 23, Table III). Nyssa talamancana thrives at middle elevations in Costa Rica and Panama under fully humid equatorial climate or fully humid warm temperate climate with warm summers (Af, Cfb; tropical-oreotropical vegetation element; File S1) together with other relict genera, e.g. Ticodendron, Alfaroa, Oremunnea and Gordonia (Hammel & Zamora 1990). Sapotaceae, which are warmth-loving and predominantly occur under equatorial (A) climates, also extend into warm temperate (e.g. Cwa, Cwb, Cfa, Cfb) climates. The cooccurrence of insect-pollinated tropical-meridional (frost-and snow-cover intolerant) and meridionalnemoral (extra-tropical) elements and the scarcity of steppe-climate (BS) tolerating taxa points towards a subtropical, humid warm temperate climate with hot summers (Cfa, Cwa) for the lowland deposition area. The taxa extending into snow (D) and polar (E) climates (Amaranthaceae, Caryophyllaceae, Andromeda, Empetrum, Ericaceae, Ludwigia) are water plants and/or herbaceous plants or small shrubs with a cosmopolitan distribution, and therefore more or less climate independent. The climatic preferences of the taxa presented herein support our previously interpreted climate signal based on the fossil Fagales to Rosales from the Sarmatian of the Lavanttal Basin (Grímsson et al. 2016, figure 23, table III). The lowlands in the Lavanttal region probably thrived in fully humid warm temperate (Cfa) climate, with possible dryer winters than summers (→ Cw climate), subtropical conditions as found today in south-eastern part of the United States and southern China (see also 23. Köppen signatures of potential modern analogues of Myrtales, Malvales, Santanales and Ericales lineages found at the Lavanttal site. The bar chart shows the proportion of extant species part of the modern genus/lineage categorised for generalised climate-vegetation types (see Denk et al. 2013;Grímsson et al. 2016; see also Material and methods section). Boreal-nemoral elements preference for D-climates and C-climates, occurring in snow and temperate climates with hot to cool summers; nemoral elements preference for warm temperate and/or snow climates with warm summers (Cfb, Cwb, Csb, Dfb, Dwb, Dsb); meridio-nemoral elements preference for warm temperate climates with hot, but not warm, summers (Cfa-and Cwa-climates); semihumid-meridional elements preference for semihumid warm temperate climates with hot (and warm) summers; tropical-meridional elements preference for tropical (A-climates) and warm temperate climates with hot but not warm summers; tropical elements species restricted to tropical (A-climates); eurytropical elements preference for non-tropical climates with summer draught and generally dry climates (B-and Cs-climates); oreotropical elements species restricted to temperate climates along altitudinal successions within the tropical zone (Cfa, Cfb, Cwa, Cwb).  Grímsson et al. 2016, figure 1). From the lowlands up into the mountains, subsequent altitudinal succession would have occurred, and the climate gradually shifted from warm temperate into snow (D) climates (Cfa → Cfb/Dfa → Dfb) providing niches for species intolerant to summer heat.
The angiosperm pollen described herein as well as palynomorphs previously presented by Grímsson et al. ( , 2015aGrímsson et al. ( , 2016a and  represent the major part of the palynoflora from the Sarmatian of the Lavanttal Basin. The remaining angiosperm (and unknown) pollen types will be described in a following contribution. A detailed interpretation and comprehensive discussion of the palaeovegetation, ecology and paleoclimate is envisaged to be presented after the remaining pollen types have been described.