Combined LM and SEM study of the Middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin: Part I. Bryophyta, Lycopodiophyta, Pteridophyta, Ginkgophyta, and Gnetophyta more

Grana, 2011; 50: 102–128 Combined LM and SEM study of the Middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin, Austria: Part I. Bryophyta, Lycopodiophyta, Pteridophyta, Ginkgophyta, and Gnetophyta FRIÐGEIR GRÍMSSON, REINHARD ZETTER & CHRISTIAN BAAL Department of Palaeontology, University of Vienna, Austria Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Abstract Preliminary studies of the palynoflora from the Lavanttal Basin show a relatively rich assemblage of pollen and spores. The palynoflora comprises at least 17 different kinds of spores, representing the Bryophyta (Sphagnum), Lycopodiophyta (Lycopodium, Selaginella), and the Pteridophyta (Dryopteris, Osmunda, Pteris), about 20 different pollen types of conifers assignable to Cupressaceae and Pinaceae, and 130–160 different kinds of angiosperm pollen. In this study, we describe all spores together with pollen from two seed plants, i.e. Ginkgo (Ginkgophyta) and Ephedra (Gnetophyta). The fossil spores and pollen grains are preserved in phosphoritic nodules. Absence of palynomorphs characteristic of marine settings and presence of numerous freshwater algae (diatoms, dinoflagellates, and different green algae) indicate freshwater environments. This is also supported by sedimentological observations suggestive of wetland surroundings, characterised by lakes, swamps, streams, rivers and floodplain areas. The taxa reported here all seem to represent part of azonal vegetation with plants growing in swamps, on hummocks, along border of lakes or streams, on levees, or on sandy patches of floodplains. Preliminary results suggest that the vegetation thrived under a relatively warm and humid climate. Keywords: Cainozoic, Carinthia, fossils, Northern Hemisphere, palynomorphs, phosphoritic nodules, pollen, spores Middle Miocene Sarmatian sedimentary deposits, c. 12.7–11.6 Ma, are found only in the Central Paratethys region (Austria, Czech Republic, Hungary, and Slovakia). The Sarmatian stage was originally defined in the Vienna Basin as a regional stage by Suess (1866; cf. Harzhauser & Piller, 2004; Piller et al., 2007) and corresponds to the Middle to Upper Serravallian (for correlation between regional and standard stages see Figure 6 later). In the Eastern Paratethys, contemporaneous sediments belong to the Volhynian and the Lower Bessarabian stages (Bulgaria, Romania, and Ukraine; cf. Harzhauser & Piller, 2004). Sarmatian sediments in the Central Paratethys are stratigraphically important as they document significant environmental changes caused by isostatic movements and intercalated transgressional and regressional facies. In the aquatic territory of the region, the invertebrate fauna reduced and became highly endemic. It is believed that this resulted from isolation of the Paratethyan Sea from the ‘Mediterranean Sea’ and the formation of intracontinental sea and lake systems (cf. Steininger & Wessely, 2000; Piller et al., 2007). Around the same time, central Europe also experienced gradual floristic turnovers in terrestrial realms caused mainly by climate cooling and increased seasonality (cf. Zachos et al., 2001; Mosbrugger et al., 2005). In recent studies by Kvaˇ ek et al. (2006), Kovar-Eder et al. (2008), c and Ivanov et al. (2011), the authors tried to reconstruct the changing vegetation and climate during the Miocene in the Central and Eastern Paratethys. Fossil floras representing the Badenian Stage, c. 16– 12.7 Ma, are apparently abundant and the same is true for the Early Pannonian Stage, starting c. 11.6 Ma ago. In the Paratethys realm, Sarmatian floras representing the time period between the Badenian and the Early Pannonian are extremely rare, with less than a handful of floras known from Serbia and Correspondence: Friðgeir Grímsson, Department of Palaeontology, University of Vienna, Althanstraße 14 (UZA II), A-1090 Vienna, Austria. E-mail: fridgeir.grimsson@univie.ac.at (Received 12 January 2011; accepted 22 April 2011) ISSN 0017-3134 print / ISSN 1651-2049 online © 2011 Collegium Palynologicum Scandinavicum DOI: 10.1080/00173134.2011.585804 Sarmatian palynoflora from Lavanttal 103 Ukraine (cf. Kovar-Eder et al., 2008; Ivanov et al., 2011). Because of the scarcity of Sarmatian floras and their absence in the western Central Paratethys, the Lavanttal palynoflora provides essential missing data for the vegetation and climate history of this time period in the region. Sarmatian floras are rare in Austria. At present, only four floras from this time interval are documented. From the Vienna Basin, Berger and Zabusch (1952, 1953) described the Türkenschanze macroflora from Döbling (18th district of Vienna), and Ettingshausen (1851), Stur (1867), and Berger (1953) published on the macroflora from Hernals (17th district of Vienna). Outside the Vienna Basin, the only Sarmatian flora that has been described so far comes from the Lavanttal Basin, Carinthia (Berger, 1955). The leaf macrofloras from these floras vary considerably in diversity, from five taxa in the Lavanttal Basin (Berger, 1955) to 50 taxa recorded for the Vienna Basin (Berger & Zabusch, 1953). An ongoing study on the Sarmatian Gratkorn flora (Styria, southeast Austria), comprising leaves, seeds, and fruits, has yielded about 25 taxa so far (Meller, 2006). Sarmatian species diversity in the Lavanttal Basin was also documented in a palynological study by Klaus (1984). This work was published in German and based mostly on light microscopy (LM) observations. The author reported a total of 27 conifer species including three species of Abies, four species of Cathaya, two species of Cedrus, one species of Keteleeria, one species of Picea, 12 species of Pinus, one species of Pseudotsuga, and three species of Tsuga. New knowledge on extant conifer pollen including information on morphological variability within single species indicates that the number of species recorded by Klaus (1984) is probably too high. Grains assigned to different species of Cathaya most likely belong to a single biological species, and some of the Pinus species should be merged. Klaus (1984) may also have overestimated the number of species for other groups. For instance, he recognised four species of Fagus, but based on variation documented for extant Fagus (cf. Denk, 2003), the morphological differences noted by Klaus (1984) probably reflect natural variability and/or different stages of maturity within a single species. In contrast, Klaus (1984) probably underestimated the species diversity for other taxa particularly among angiosperms. We have, therefore, undertaken a combined LM and scanning electron microscopy (SEM) study of the Sarmatian palynoflora from the Lavanttal Basin (Figures 1, 2) to obtain a higher and more reliable taxonomic resolution. This study will also result in a more accurate basis for accessing vegetation composition and species diversity of the palaeoflora and, therefore, improve palaeoenvironmental interpretations and reconstruction of the vegetation for this time in the western Central Paratethys. Due to the high species diversity, our results will be published in a series of five papers. The first part provides information on material and methods used, and a thorough overview of geology and sedimentary environments. The taxonomic section includes descriptions of bryophyte, lycophyte, and pteridophyte spores together with pollen of Ginkgo and Ephedra. The second part will treat the coniferous component, and the third and fourth parts will describe the angiosperm pollen. Finally, the fifth part will deal with the palaeoenvironment, composition of the flora, and comparison with coeval neighbouring floras, vegetation types, as well as climate signals. This study is the first to describe an entire palynoflora based consistently on LM and SEM to identify palynomorphs. This allows for much higher taxonomic resolution than classical palynological studies using LM only, and it is our hope that the study may serve as a reference flora for the Middle Miocene of Central Europe. Materials and methods Preparation of samples Phosphoritic nodules were washed and dried and then placed into small glass beakers filled with 350 ml of concentrated hydrochloridic acid (HCl) for one day. Nodules were fully dissolved after this time. After decanting most of the HCl residue, the remaining substance was boiled for c. ten minutes in 200 ml hydrofluoric acid (HF). The solution was then transferred into a four litre beaker filled with water. After settling, the liquid was decanted and the remaining substance was again boiled in HCl for five minutes. After cooling and settling, most of the HCl was decanted and the remaining solution was centrifuged; the deposit was then washed three to four times with water. The sample was acetolysed, washed again with water, and centrifuged up to four times. The final remaining organic material was then mixed with glycerol and stored in small sample tubes. Preservation of palynomorphs The samples resulting from the prepared nodules (Figure 3) are rich in organic material (Figure 5A). The preservation is extremely good, the palynomorphs are preserved in a three-dimensional (3D) state and look almost like extant material. The excellent preservation is due to the fact that the pollen and spores are preserved in phosphoritic Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 104 F. Grímsson et al. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 1. A–C. Maps showing the geographical position of the Lavanttal Basin in Europe and major geological formations. D. Detailed geological map of the area surrounding the basin (modified after Egger et al., 1999; Bechtel et al., 2007; Reischenbacher et al., 2007); the position of the Lavanttal Basin is marked with a broken line. Sarmatian palynoflora from Lavanttal 105 Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 2. Schematic cross-sections showing the sedimentary units filling the basin and simplified geological map of the Lavanttal Basin. Area in box enlarged from Figure 1D (modified after Beck-Mannagetta, 1952; Bechtel et al., 2007; Reischenbacher et al., 2007). The nodules used in this study have been collected in an underground mine at St Stefan; for stratigraphic position of the nodules see uppermost cross-section. 106 F. Grímsson et al. nodules/concretions. During fossilisation, which must have happened relatively soon after deposition, the palynomorphs were not compacted and mostly kept or at least regained their optimal 3D form and become close to fully hydrated after treatment (see picked grains in Figure 5B). Further, the palynomorphs were apparently not affected by any oxidisation process or microbial destruction, making all surface features perfectly visible under SEM. However, many of the palynomorphs contained small pyrite (FeS2 ) crystals inside their cavities. Description of taxa All descriptions of spores and pollen include the most diagnostic features observed in LM and SEM. Terminology for description follows mostly Punt et al. (2007) for LM, and Hesse et al. (2009) for SEM. The systematic section starts with Bryophyta (mosses), Lycopodiophyta (clubmosses), and Pteridophyta (horsetails and true ferns), followed by the Ginkgophyta and the Gnetophyta. Families and genera appear in alphabetical order. Incertae sedis are listed at the end of each larger taxonomic group. Conservation of material Parts of all the original sedimentary rock samples (the nodules), all treated samples stored in tubes, and SEM stubs of fossil palynomorphs are stored in the collection of the Department of Palaeontology, University of Vienna, under accession numbers 5150/01-95. The single grain technique Fossil palynomorphs were investigated both by LM and SEM using the single grain technique (cf. Zetter, 1989; Hesse et al., 2009). This technique has already proven to be very useful when studying fossil palynofloras, compared to using LM only, when a detailed and more accurate systematic identification of the spores and pollen are needed (cf. Ferguson et al., 2007; Grímsson et al., 2008; Denk et al., 2010), and resulting in an higher taxonomic resolution. For preparation, drops from the sample tubes were transferred onto LM slides and solitary grains were picked using a preparation needle with a single hair mounted on it. The grains were placed on a separate slide with clean glycerol for study and photography in LM. The same grains were then transferred on a SEM stub with the help of the preparation needle and washed with drops of absolute ethanol to dissolve the remaining glycerol. The stubs with the palynomorphs were then sputter coated with gold and the objects studied and photographed under SEM. Using this method, the harmomegathic effect (cf. Hesse et al., 2009), which relates to the dry or hydrated state of the pollen grain, sometimes becomes obvious. In glycerol, the pollen grains can become as hydrated as possible and obtain their ‘optimal’ form. When the pollen grains are transferred over to the SEM stubs, they sometimes react to the absolute ethanol by drying up and shrinking. The pollen can become between 5–30% smaller in SEM than in LM. This depends on the pollen wall structure, thickness of the wall, and the size of the pollen grain. The highest shrinking is noticed in pollen grains equipped with air sacks such as pollen of the Pinaceae. The shape of the pollen can also be different in SEM and in LM after treating with ethanol. The more dried pollen grain can become oblate in SEM but have been prolate or spheroidal in LM and vice versa. Large pollen grains with a relatively thin wall compared to size can collapse and become infolded in various ways. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Geological background Geological setting and age of the sediments The geological history of Austria is complex and the geology around the study area highly variable (Figure 1). For this reason, we refer only to the basic geological outline and time period necessary. For a more detailed view, we refer to Beck-Mannagetta (1952), Bassir (1964), Janoschek and Matura (1980), Oberhauser (1980), Weber and Weiss (1983), and Krenmayr et al. (2000). The evolution of the Eastern Alps during the Early and Middle Miocene was controlled by tectonic extension and lateral extrusions (cf. Ratschbacher et al., 1991; Frisch et al., 2000), as well as the formation of many intramontane basins (cf. Márton et al., 2000). The sinistral northern border of the main extruding block was located in the Northern (Mesozoic) Calcareous Alps, whereas the dextral Peri-adriatic Lineament outlined its southern border (Figure 1B, C). During the Early Miocene, several basins started to subside almost synchronously on top of the extruding block of the easternmost Alps. These include the Styrian Basin, the Vienna Basin, and the basins along the Mur-Mürz Fault System and the Lavanttal Fault System (Peresson & Decker, 1997; Haas et al., 1998). In some areas, the subsidence and resulting sedimentation continued well into the Late Miocene (Figure 6; Márton et al., 2000; Harzhauser & Piller, 2004; Reischenbacher et al., 2007). Several Miocene basins formed along the Lavanttal Fault System, among them the Lavanttal Basin. Today, the northwest (NW)-southeast (SE) Sarmatian palynoflora from Lavanttal 107 Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 3. Phosphoritic nodules. A–C. Nodules preserved within a thin erosive conglomerate, allochthonous. D. Various shaped nodules freed from their surrounding sediments. E–F. Nodule preserved within a fine grained diatom rich marlstone, autochthonous. a – small stones and pebbles within the conglomerate; b – transported and coalified wood fragments; c – traces left by removed nodules; d – phosphoritic nodules. Scale bar − 5 cm. Figure 4. One of the original labels from Beck-Mannagetta’s phosphoritic nodule collection dating back to the year 1949. trending Lavanttal Fault System is still active and dextrally offsets the Peri-adriatic Lineament for 10 to 14 km (Vrabec et al., 2006). The sedimentary succession and stratigraphic division of the Lavanttal Basin (Figures 1D, 2) has been presented by Beck-Mannagetta (1952) and further by Bassir (1964), Bechtel et al. (2007), and Reischenbacher et al. (2007). The oldest sediments are fluvial deposits belonging to the Granitztal Formation. These late Lower Miocene (Carpatian) deposits overlie the much older crystalline basement rock and the adjacent Mesozoic sediments (Figure 2). The Granitztal Formation is, in turn, overlain by the Mühldorf Formation that is subdivided into a lower lacustrine part (fish shale) and a following marine sequence. The Mühldorf Formation has been considered to be of early Middle Miocene (Early Badenian) age, based on 108 F. Grímsson et al. Figure 5. Pollen isolation. A. Organic material as seen in the glycerol, including amongst others various algae, pollen, spores, and cuticle pieces. B. Pollen grains picked from the main sample such as in (A) and transferred to a fresh drop of glycerol. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 both biostratigraphic correlations of various fossils (Beck-Mannagetta, 1952; Weinfurter, 1952; Berger, 1955; Kühn, 1963; Schmid, 1974, Meller & Kvaˇ ek, c 2007; Reischenbacher et al., 2007) and absolute age determinations of c. 14.9 Ma (Lippolt et al., 1975) from the Kollnitz basalts and associated tuffs within the formation. Most recent chronostratigraphic and biostratigraphic correlations based on calcareous nannoplankton, ostracods, and foraminifera give an age of 14.9 to 14.7 Ma for the upper marine part of the Mühldorf Formation (Reischenbacher et al., 2007). Sediments above the Mühldorf Formation are of marine origin, but fossils reported from these sediments suggest a gradual transition to brackish conditions during the Middle Badenian to pure freshwater conditions in the Late Badenian (cf. Beck-Mannagetta, 1952). Brackish and freshwater deposits continued to accumulate during the Early Sarmatian in the central part of the Lavanttal Basin (Figure 2). The freshwater sediments comprise three coal seams (Beck-Mannagetta, 1952; Bechtel et al., 2007; Reischenbacher et al., 2007) and, above the uppermost coal seam, phosphoritic nodules are embedded within a diatomite-rich fine-grained marlstone (Figure 2). These phosphoritic nodules contain the palynomorphs described in this study. The subsequent sediments are also of freshwater origin and of Late Sarmatian to Early Pannonian age (cf. BeckMannagetta, 1952). The gravels, sandstones (sand), and diatom rich marlstones (including the coal seams) from the Lower Sarmatian of the Lavanttal Basin (Figure 2) correlate well with other coeval lithological units from, for example, the nearby Styrian Basin (Figure 6; cf. Harzhauser & Piller, 2004) that are all included into the Lower Sarmatian fourth order LS-1 sequence [according to the sequence stratigraphy by Harzhauser and Piller (2004)], which started c. 12.7 Ma and ended c. 12.1 Ma. The brackish sediments on top of the coal seams and the phosporitic nodules are here believed to represent the initial phases of the Lower Sarmatian maximum flooding event in the western Central Paratethys. This event corresponds with the maximum transgression during the so-called Mohrensternia Zone that lasted from c. 12.7 Ma until c. 12.4 Ma (Figure 6; cf. Harzhauser & Piller, 2004). A more precise age of the nodules is hard to pinpoint, but given their stratigraphical position within the sequence, the thickness of the sediments surrounding the coals seems, estimated accumulation rates, and hiatus events above the seams, they are most likely between 12.6 and 12.5 Ma. The phosphoritic nodules and the sedimentary environment The phosphoritic nodules (Figure 3A–F) have been part of the collection of the Department of Palaeontology of the Vienna University for over 60 years. Most of the nodules and sedimentary boulders containing nodules were collected by Dr BeckMannagetta or given to him by his acquaintances (cf. Klaus, 1984). The nodules were collected during the late 1940s until the late 1960s (Figure 4). The nodules are preserved in diatom rich marlstones just above the uppermost Lower Sarmatian coal seam (Figure 2). They have a rather restricted distribution but where they occur, they do so in high numbers (Beck-Mannagetta, 1952; Bassir, 1964). From the underground mine of St Stefan, Bassir (1964) reported more than 100 nodules per square metre of sedimentary surface directly above the coal seam. Close examination reveals that the sediment comprising the numerous densely packed nodules is a relatively thin erosive conglomerate horizon Sarmatian palynoflora from Lavanttal 109 Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 6. Stratigrapy of the Sarmatian sedimentary basins in Austria (modified after Harzhauser & Piller, 2004; based on data from: Grill, 1941; Papp et al., 1974; Haq et al., 1988; Cicha et al., 1998; Hardenbol et al., 1998; Hilgen et al., 2000; Lirer et al., 2002; Harzhauser & Piller, 2004). within the fine-grained laminated marlstone unit (Figure 3A–C). Here, the nodules are assumed to be allochthonous. Nodules that are preserved in marlstone (Figure 3E, F) outside the conglomerate are rare and assumed to be autochthonous (Bassir, 1964). From the sedimentary structures around the individual nodules, it appears that the nodules were increasing in size within the sediment or at the sediment–water interface during continuous sedimentation before the sediments became lithified or fossilised. The nodules are spherical to oblate in shape, sometimes occurring as twins. The surface of the nodules is smooth, shiny, and of greyish to blackish colour (Figure 3D). In crosssection, the darker margin measures 0.5–1.5 mm and is marked by a sudden but uneven change from the more greyish to brownish mass composing most of the nodule. Without magnification and with a non-polished surface, the nodule displays no growth-rings or radiating striation that would suggest any periodical or seasonal growth. The nodules have distinct inorganic nuclei of variable sizes from where the phosphatic precipitation commenced. These nuclei are not only found in the centre of the nodules but also close to the margin (Figure 3D). Analyses of the nodules by Bassir (1964) have shown that the nodules are composed of c. 90% tri-calcium phosphate [Ca3 (PO4 )2 ] and 10% clay. The phosphorous pentoxide (P2 O5 ) content is approximately 40%. 110 F. Grímsson et al. The formation of phosphorite (phosphatic sediments) and, particularly phosphoritic nodules, has been debated for years. Phosphoritic sediments, sedimentary rocks, and nodules have been found in many parts of the world, in recent environments as well as in localities representing different time periods of the geological record. Based on these findings, many theories concerning the formation of the nodules, the origin of the phosphate, if it is organic or inorganic, if it is primary or secondary, and what were the controlling factors causing the accumulation, have been put forward (cf. Soudry & Lewy, 1988; Reimers et al., 1990; Lucas & Prévôt, 1991; Ogihara, 1999; Schmid, 2000). All fossil phosphoritic nodules from the geological record have until now been reported from marine or brackish sediments. In contrast, the nodules from the Lavanttal Basin were definitely formed in a freshwater environment because in the fine-grained sediments surrounding the nodules and inside the nodules, marine palynomorphs and marine macrofossils are absent, and only large amounts of freshwater algae and terrestrial palynomorphs are preserved. There are relatively few species of freshwater algae present but most of them occur in high numbers. So far, we have been able to identify different freshwater diatoms, various green algae, and freshwater dinoflagellates. The diatoms comprise about four to five species with a single dominant type belonging to the phytoplanktonic genus Aulacoseira (Aulacoseiraceae). Diatom rich samples are full with various structural parts from this type (Figure 7). The members of this genus are well known from recent freshwater environments (Hustedt, 1942; Krammer & Lange-Bertalot, 1991; Brian et al., 2002; Wehr & Sheath, 2003) and have been reported from many sedimentary rocks of lacustrine origin and, therefore, indicate freshwater environments (Hayworth & Sabater, 1993; ChaconBaca et al., 2002; Ambwani et al., 2003; Wolfe & Edlund, 2005; Wolfe et al., 2006; Houk, 2007). The samples all contain a high number of Botryococcus colonies (Botryococcus cf. braunii Kützing; Figure 8A–C). Even though extant Botryococcus can sometimes be found in brackish waters, it generally lives in freshwater swamps, pools, ponds, and lakes (Batten & Grenfell, 1996). Botryococcus has a long fossil record and algal colonies assigned to the genus have been reported throughout the whole Cainozoic as well as the Mesozoic (cf. Batten & Grenfell, 1996). Freshwater green algal colonies of Pediastrum are also numerous in all the samples. Until now, we have identified two different species, Pediastrum boryanum (Turpin) Meneghini (Figure 8D–F) and Pediastrum duplex Meyen (Figure 8G–L). Extant and subfossil Pediastrum algae are known to inhabit freshwater environments and are often used for environmental and climatic reconstructions of palaeo-lakes and their surroundings (cf. Parra Barrientos, 1979; Komárek & Jankovská, 2001). Pediastrum algae are also well known from the fossil record of freshwater sediments in Austria (Zetter, 1987). The dinoflagellates in the Lavanttal samples are comparable to other fossil freshwater dinoflagellates belonging to the genera Messelodinium, Geiselodinium, and Pseudokomewuia. Dinoflagellates of these types were widely distributed throughout the Cainozoic of Europe and North America (Krutzsch, 1962; Krutzsch & Pacltová, 1990; Batten et al., 1999; Lenz et al., 2007a, 2007b). Systematic palaeontology Division Bryophyta Family Sphagnaceae Dumortier Genus Sphagnum L. Sphagnum sp. (Figure 9A–C) Description. — Spore, monad, trilete, shape oblate, convex-triangular in polar view, elliptic in equatorial view; equatorial diameter 26–29 µm in LM, 22–24 µm in SEM, polar axis 14–15 µm long in LM; laesurae 2/3 of the spore radius, laesurae slightly protruding (SEM); exospore 1.3–1.8 µm thick, inner exospore thicker than outer exospore, apices slightly thickened (LM), distal thickening reaching into proximal face (SEM); sculpturing of proximal face more or less psilate, distal face verrucate (SEM). Remarks. — Fossil Sphagnum spores similar to the Lavanttal type have often been described under the fossil genera Distancoraesporis and/or Stereisporites (cf. Stuchlik et al., 2001). Ecological implications. — There are between 100–300 Sphagnum species worldwide, most of them occurring in the Northern Hemisphere (cf. Buck & Shaw, 2001). Sphagnum mosses are common in most wetland environments, thriving in and around swamps, fens, bogs, ponds, and lakes in cold tundra to warm temperate climates (cf. Rydin et al., 2006a). Division Lycopodiophyta Family Lycopodiaceae Mirbel Genus Lycopodium L. Lycopodium sp. 1 (Figure 9D–F) Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Sarmatian palynoflora from Lavanttal 111 Description. — Spore, monad, trilete, shape oblate, outline convex-triangular in polar view, elliptic in equatorial view, equatorial diameter 31–36 µm in LM, 29–32 µm in SEM; exospore 3.7–4.1 µm thick (LM); sculpturing of distal face reticulate, muri of variable width, diameter of brochi varying (SEM). Remarks. — Spores of this type and Lycopodium sp. 2 (see later) have often been described within the fossil genus Retitriletes (cf. Stuchlik et al., 2001). Ecological implications. — The genus Lycopodium comprises c. 40 extant species with a cosmopolitan distribution (Øllgaard, 1987). In North America, Lycopodium is distributed from the far north southwards to the northern border of Alabama, Georgia, and South Carolina in the southeast USA, thriving in cold tundra to warm temperate climates. The plants also grow in various habitats and occur in open grassy fields, broadleaved forests, swampy or moist conifer forests, mountain forests, or rocky outcrops, ranging in altitude from sea level up to c. 1900 m (Flora of North America Editorial Committee, 1993). From Malaysia, Rusea et al. (2009) reported Lycopodium from the understory of lowland to upper mountain forests where it occurred mostly in open sunny patches within woodland vegetation. Lycopodium sp. 2 (Figure 9G–I) Description. — Spore, monad, trilete, shape oblate, outline convex-triangular in polar view, elliptic in equatorial view, equatorial diameter 40–42 µm in LM, 32–35 µm in SEM, polar axis 34–36 µm long in LM; laesurae > 2/3 of the spore radius, laesurae slightly protruding (SEM); exospore 3.5– 4.5 µm thick; sculpturing of distal face reticulate, muri consistent in width and straight, diameter of brochi relatively constant, proximal face incomplete reticulate, showing low relief muri (SEM). Family Selaginellaceae Willkomm Genus Selaginella Palisot de Beauvois Selaginella sp. (Figure 9J–L) Description. — Spore, monad, trilete, shape oblate, circular to convex-triangular in polar view, elliptic in equatorial view, equatorial diameter without echinae 24–27 µm in LM, 19–21 µm in SEM, polar axis 16–18 µm in LM; laesurae 2/3 of the spore radius, Description. — Spore, monad, monolete, shape oblate, elliptic in polar view, elliptic to kidney formed in equatorial view, equatorial diameter 36–43 µm in LM, 30–38 µm in SEM, polar axis 19–31 µm in LM, 19–25 µm in SEM; laesura 21–30 µm long in LM, 17–25 µm in SEM; exospore 0.9–1.2 µm thick, inner exospore thinner than outer exospore; sculpturing of exospore psilate, the perispore is rugulate-cristate when preserved (SEM). laesurae indistinct (LM, SEM); exospore 1.8–2.0 thick, inner exospore as thick as outer exospore (LM), distal thickening reaching into proximal face (SEM); sculpturing echinate, distal face with long echinae and perforations, in proximal face echinae become shorter towards centre, central part of proximal face with fossulae and verrucae (SEM). Remarks. — This spore type is comparable with spores from extant species of both the Subgenus Selaginella and Subgenus Stachygynandrum as described by Tryon and Lugardon (1991), but both subgenera compose spores with relatively long echinae. Fossil spores of this type have often been described within the fossil genus Echinatisporis (cf. Stuchlik et al., 2001). Ecological implications. — The subgenus Selaginella is composed of only two extant species, S. selaginoides (L.) Beauv. Ex Mart. et Schrank that has a more or less circumboreal Northern Hemispheric distribution, and S. deflaxa Brack. that is native to Hawaii (cf. Tryon & Lugardon, 1991; Flora of North America Editorial Committee, 1993). Selaginella selaginoides often occurs in wetlands, in close vicinity to swamps, along lake margins and streams, on wet rock substrate, but mostly occurs in an open vegetation cover (cf. Flora of North America Editorial Committee, 1993). The subgenus Stachygynandrum is composed of c. 600 extant species with a subtropical to tropical circumpolar distribution. Many of the species thrive in lowlands and at moderate elevation along riverbanks, lake margins, swamps, and in moist ravines, but some also grow on drier substrates such as hummocks or rocky outcrops. The latter is very species dependent (cf. Kramer & Green, 1990; Flora of North America Editorial Committee, 1993). Division Pteridophyta Family Dryopteridaceae Herter Genus Dryopteris Adanson Dryopteris sp. (Figure 10A–F) Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 112 F. Grímsson et al. Remarks. — Spores of this type are comparable with extant Dryopteris taxa as described by Tryon and Lugardon (1991). Similar fossil spores without perispore have been described belonging to the fossil genus Laevigatosporites (cf. Stuchlik et al., 2001). Ecological implications. — The genus Dryopteris is composed of c. 225 extant species with a subcosmopolitan distribution in temperate to tropical environments (cf. Kramer & Green, 1990). Dryopteris occurs in wetland environments along margins of swamps and banks of streams and on dryer substrate such as hummocks and even on rock walls. The plants are also thriving as undergrowth in moist rocky woodlands of lowland and foothill forests and in ravines, with their distribution extending up to high elevations on mountains and even alpine forests (cf. Kramer & Green, 1990; cf. Flora of North America Editorial Committee, 1993). Family Polypodiaceae Bercht. et J.S. Presl Polypodiaceae gen. et spec. indet. 1 (Figure 10G–I) Description. — Spore, monad, monolete, shape oblate, elliptic in polar and equatorial view; equatorial diameter 50–52 µm in LM, 47–49 µm in SEM, polar axis 30–32 µm in LM, 31–33 µm in SEM; laesura 30–33 µm long in LM, 26–28 µm in SEM; exospore 1.4–1.7 µm thick, exospore thinner in central proximal face; sculpturing of exospore verrucate, verrucae widely spaced and becoming small towards central part of proximal face (LM, SEM). Remarks. — Extant spores comparable with this fossil spore type and the Polypodiaceae gen. et spec. indet. 2 (see later) are mostly found within the family Polypodiaceae, but also within the family Davalliaceae (Tryon & Lugardon, 1991). Fossil spores similar to these Lavanttal spore types have been included in the fossil genus Verrucatosporites (cf. Stuchlik et al., 2001). Polypodiaceae gen. et spec. indet. 2 (Figure 10J–L) Description. — Spore, monad, monolete, shape oblate, elliptic in polar view, elliptic to kidney-shaped in equatorial view; equatorial diameter 71–74 µm in LM, polar axis 46–55 in LM; laesura 39–41 µm long in LM; exospore 3.3–3.5 µm thick, distal thickening reaching into proximal face, exospore thinner in central proximal face (LM); sculpturing verrucate, verrucae densely packed and becoming smaller towards central part of proximal face. Family Pteridaceae Reichenbach Genus Pteris L. Pteris sp. 1 (Figure 12A–C) Description. — Spore, monad, trilete, shape oblate, triangular in polar view, elliptic in equatorial view; equatorial diameter including cingulum 32–35 µm in LM, 32–34 µm in SEM, polar axis 23–25 µm in LM; laesurae 3/4–4/5 of the spore radius; exospore 1.0–1.5 µm thick (LM), cingulum 3–5 µm wide (SEM); sculpturing rugulate on proximal and distal face, rugulae smooth and of low relief, consistent in size, cingulum is psilate. Remarks. — Extant Pteris spores comparable with the fossil spore types Pteris sp. 1, Pteris sp. 2, and Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Family Osmundaceae Bercht. et J.S. Presl Genus Osmunda L. Osmunda sp. (Figure 11A–F) Description. — Spore, monad, trilete, shape spheroidal, circular in polar and equatorial view, equatorial diameter 48–58 µm in LM, 41–52 µm in SEM; laesurae 1/2–2/3 of the spore radius, laesurae protruding (SEM); exospore 2.8–3.1 µm thick (LM); sculpturing rugulate (tuberculate), echinate to microechinate (SEM). Remarks. — The considerable size range noted in the fossil material is also observed in studies on extant Osmunda species (cf. Tryon & Lugardon, 1991). Fossil spores comparable to the Lavanttal Osmunda type have often been described within the fossil genus Baculatisporites (cf. Stuchlik et al., 2001). Ecological implications. — Osmunda is a small genus composed of only 6–8 extant species that is found in most temperate to tropical regions of the world (cf. Kramer & Green, 1990; Metzgar et al., 2008). Osmunda generally occurs in major wetlands of lowland areas in or at the margin of marshes, fens, swamps, lakes, and streams. It is also found growing in ponds and along streams in deciduous or coniferous upland forests, and Osmunda regalis L. apparently typifies small ponds throughout the temperate forest regions in North America and Europe (cf. Flinn et al., 2008; Landi & Angiolini, 2011). Sarmatian palynoflora from Lavanttal 113 Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 7. SEM micrographs of the fossil freshwater diatom Aulacoseira from the phosphoritic nodules. A. Oblique valvular view with poroids. B–F. Cingular views: B. With cingulun; C. Linkage of adjacent cells by connecting spines; D. Separation valve with long spines; E. Parts of cingulum; F. Without cingulum. G. Oblique valvular view, small rimoportulae inward of the ‘Ringleiste’. H, I. Cingular views: H. Covered by cingulum; I. Linkage of adjacent cells by connecting spines. J–L. Details: J. Rimoportulae and Ringleiste; K. Valve, mantle areolae with volate occlusions; L. Valve, mantle areolae and ligula of cingulum. Scale bars − 10 µm (A–I), 1 µm (J–L). 114 F. Grímsson et al. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 8. LM (A, D, G, J) and SEM (B, C, E, F, H, I, K, L) micrographs of fossil freshwater algae from the phosphoritic nodules. A–C. Botryococcus cf. braunii. D–F. Pediastrum boryanum. G–L. Pediastrum duplex. Scale bars − 10 µm (A, B, D, E, G, H, J, K), 1 µm (C, F, I, L). Sarmatian palynoflora from Lavanttal 115 Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 9. LM (A, D, G, J) and SEM (B, C, E, F, H, I, K, L) micrographs of dispersed fossil spores. A–C. Sphagnum sp. D–F. Lycopodium sp. 1. G–I. Lycopodium sp. 2. J–L. Selaginella sp. Scale bars − 10 µm (A, B, D–H, J, K), 1 µm (C, I, L). 116 F. Grímsson et al. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 10. LM (A, D, G, J–L) and SEM (B, C, E, F, H, I) micrographs of dispersed fossil spores. A–F. Dryopteris sp. G–I. Polypodiaceae gen. et spec. indet. 1. J–L. Polypodiaceae gen. et spec. indet. 2. Scale bars − 10 µm (A, B, D, E, G, H, J, K, L), 1 µm (C, F, I). Sarmatian palynoflora from Lavanttal 117 Pteris sp. 3 (see later) have been described by Tryon and Lugardon (1991). Similar fossil spores have been described within the fossil genus Polypodiaceoisporites (cf. Stuchlik et al., 2001). Ecological implications. — This is a moderately sized genus composing 200–250 extant species with a cosmopolitan distribution and occurring in warmtemperate to tropical regions (cf. Kramer & Green, 1990; Tryon & Lugardon, 1991). The plants grow mostly in moist environments and are absent in dry or temporarily dry regions. They often occur in relatively shady places as part of the undergrowth and in various forests types, with representatives at sea level as well as in high mountains. Plants are often growing in open patches within woodland and at the forest borderland. Some few species are found on a dryer rocky substrate (cf. Kramer & Green, 1990; Tryon & Lugardon, 1991). Pteris sp. 2 (Figure 12D–F) Description. — Spore, monad, trilete, shape oblate, triangular in polar view, elliptic in equatorial view; equatorial diameter including cingulum 35–37 µm in LM, 32–34 µm in SEM, polar axis 26–28 µm in LM; laesurae 1/2–3/4 of the spore radius; exospore 1.2–2.2 µm thick, cingulum 4.4–4.7 µm wide in LM, 4.7–4.8 µm in SEM; sculpturing rugulate on distal face, rugulae high and crested, varying in size and shape, rugulae decreasing in size on proximal face, cingulum is psilate (SEM). Pteris sp. 3 (Figure 12G–I) Description. — Spore, monad, trilete, shape oblate, triangular in polar view, elliptic in equatorial view; equatorial diameter including cingulum 26–28 µm in LM, 23–26 µm in SEM, polar axis 18–19 µm in LM; laesurae 2/3–3/4 of the spore radius; exospore 1.4–1.6 µm thick, cingulum 2.9–3.1 µm wide in LM, 2.5–2.8 µm in SEM; sculpturing rugulate and perforate on both distal and proximal faces, sculpturing showing lower relief in proximal face (SEM). Incertae sedis — unassigned pteridophyte spores Pteridophyta fam., gen. et spec. indet. 1 (Figure 11G–L) Description. — Spore, monad, monolete, shape oblate, elliptic in polar view, elliptic to kidney-shaped in equatorial view, equatorial diameter 40–46 µm in LM, 31–40 µm in SEM, polar axis 28–32 µm in Pteridophyta fam. gen. et spec. indet. 4 (Figure 13D–F) Description. — Spore, monad, trilete, shape oblate, triangular in polar view, elliptic in equatorial view, equatorial diameter 34–36 µm in LM, 30–31 µm in SEM; laesurae 2/3 of the spore radius (LM); exospore 0.8–1.4 µm thick (LM); sculpturing low LM, 24–30 µm in SEM; laesura 27–33 µm long in LM, 22–29 µm in SEM; exospore 1.3–1.7 µm thick, inner exospore thinner than outer exospore, exospore slightly protruding along the laesura; sculpturing microverrucate to microrugulate, perforate (SEM). Remarks. — Several extant spore types found in various families within the Pteridophyta (Tryon & Lugardon, 1991) are comparable with this fossil spore type and makes further identification impractical. Pteridophyta fam., gen. et spec. indet. 2 (Figure 12J–L) Description. — Spore, monad, trilete, shape oblate, triangular to concave-triangular in polar view, elliptic in equatorial view, equatorial diameter 31–34 µm in LM, 32–33 µm in SEM, polar axis 29–31 µm in LM, 24–28 µm in SEM; laesurae 4/5 of the spore radius, exospore 0.8–1.1 µm thick; sculpturing psilate, perforate, perforations irregularly distributed. Remarks. — Extant spore types that are similar to Pteridophyta fam., gen. et spec. indet. 2, Pteridophyta fam., gen. et spec. indet. 3, and Pteridophyta fam. gen., et spec. indet. 4 (see later) are found in various families within the Pteridophyta (Tryon & Lugardon, 1991). This makes further affiliation impractical. Comparable fossil spores from other palynofloras have been included in the fossil genus Leiotriletes (cf. Stuchlik et al., 2001). Pteridophyta fam., gen. et spec. indet. 3 (Figure 13A–C) Description. — Spore, monad, trilete, shape oblate, triangular in polar view, elliptic in equatorial view, equatorial diameter 33–36 µm in LM, 30–32 µm in SEM; laesurae 3/4 of the spore radius, laesura with curvatura imperfecta, exospore protruding over laesura (SEM); exospore 1.0–1.3 µm thick (LM), distal thickening reaching into proximal face (LM, SEM); sculpturing psilate to granulate (SEM). Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 118 F. Grímsson et al. relief microrugulate to microverrucate, perforate, perforation irregularly distributed (SEM). Pteridophyta fam., gen. et spec. indet. 5 (Figure 13G–L) Description. — Spore, monad, trilete, shape oblate, triangular to circular in polar view, elliptic in equatorial view, equatorial diameter 29–40 µm in LM, 27– 34 µm in SEM; laesurae 1/3–1/2 of the spore radius (LM); exospore 0.9–1.1 µm thick (LM); sculpturing foveolate, granulate (SEM). Remarks. — We were not able to convincingly correlate this spore type and the Pteridophyta fam. gen. et spec. indet. 6 spore type (see later) to any extant subgroup, and tentatively assigned them to the Pteridophyta until further information suggests otherwise. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Pteridophyta fam., gen. et spec. indet. 6 (Figure 14A–F) Description. — Spore, monad, trilete, shape oblate, triangular to circular in polar view, elliptic in equatorial view, equatorial diameter 32–37 µm in LM, 23–29 µm in SEM, polar axis 16–20 µm in LM; laesurae 3/4 of the spore radius (LM), exospore protruding over laesura (SEM); exospore 2.0–2.2 µm thick (LM), distal thickening reaching into proximal face (LM, SEM); sculpturing microverrucate, rugulate, fossulate on distal face, sculpturing elements decreasing is size and of lower relief in proximal face (SEM). Division Ginkgophyta Family Ginkgoaceae Engler Genus Ginkgo L. Ginkgo sp. (Figures 14G–L, 15A–C) Description. — Pollen, monad, sulcate, shape oblate (boat-shaped), elliptic in polar and equatorial views, equatorial diameter 31–38 µm in LM, 32–36 µm in SEM, polar axis 24–26 µm in LM; exine 0.9– 1.1 µm thick, nexine thinner than sexine (LM); sculpturing rugulate to microrugulate, rugulae crested (SEM). Remarks. — The characteristically boat-shaped (dry, without cell content), sulcate pollen grains with rugulate to microrugulate sculpturing are associated with the genus Ginkgo (cf. Halbritter, 2000; Zhang et al., 2000; Nakao et al., 2001). This fossil pollen type fits well within the morphological range known from pollen of the extant Ginkgo biloba L., but it is unlikely that this fossil type belongs to the extant species. The pollen record of Ginkgo is relatively poor (Zhou, 2009) as most palynological studies are based on LM observations only. Under LM it is hard to distinguish between real Ginkgo pollen and various Cycadaceae pollen grains. However, with combined LM and SEM investigations of Cainozoic palynofloras from Eurasia by Gastaldo et al. (1998) and Hofmann and Zetter (2005), and from North America by Zetter et al. (in press), the authors frequently encountered Ginkgo pollen. The macrofossil record of the genus Ginkgo has shown that Late Cainozoic European findings all belong to the same fossil taxon Ginkgo adiantoides (Ung.) Heer (based on leaf remains), whereas from Asia, two co-occurring Ginkgo species, i.e. G. adiantoides and G. jiayiensis C. Quan, G. Sun et Z. Zhou are known (cf. Denk & Velitzelos, 2002; Quan et al., 2010). It is quite plausible that the fossil Ginkgo pollen from the Lavanttal Basin belong to the same biological species as the widespread Late Cainozoic European G. adiantoides leaf remains. Ecological implications. — Ginkgo is a relict genus comprising only the single living species G. biloba. Ginkgo trees are deciduous and known only from a restricted area in east China, where they compose merely a minor part of the broadleaved forests. The trees grow mostly in lowland regions, valleys, and on mountain slopes, at elevation between 300–1100 m, along the Yangtze River (Flora of China Editorial Committee, 1999). According to del Tredici et al. (1992), Ginkgo grows mostly in disturbed microsites, along streams and rivers, on steep rocky substrates, and at the edge of cliffs. It has also been suggested that the trees are gap opportunists and can cope well as an understory element under shady conditions until they get the chance to become a canopy tree when a gap occurs (del Tredici, 1989). Royer et al. (2003) concluded that Late Cretaceous–Middle Miocene Ginkgo trees were mostly confined to disturbed areas, growing along streams and rivers and in levée environments. Division Gnetophyta Family Ephedraceae Dumortier Genus Ephedra L. Ephedra sp. 1 (Figure 15D–I) Description. — Pollen, monad, inaperturate, oblate, elliptic in polar view, star-like in equatorial view Sarmatian palynoflora from Lavanttal 119 Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 11. LM (A, D, G, J) and SEM (B, C, E, F, H, I, K, L) micrographs of dispersed fossil spores. A–F. Osmunda sp. G–L. Pteridophyta fam., gen. et spec. indet. 1. Scale bars − 10 µm (A, B, D, E, G, H, J, K), 1 µm (C, F, I, L). 120 F. Grímsson et al. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 12. LM (A, D, G, J) and SEM (B, C, E, F, H, I, K, L) micrographs of dispersed fossil spores. A–C. Pteris sp. 1. D–F. Pteris sp. 2. G–I. Pteris sp. 3. J–L. Pteridophyta fam., gen. et spec. indet. 2. Scale bars − 10 µm (A, B, D, E, G, H, J, K), 1 µm (C, F, I, L). Sarmatian palynoflora from Lavanttal 121 Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 13. LM (A, D, G, J) and SEM (B, C, E, F, H, I, K, L) micrographs of dispersed fossil spores. A–C. Pteridophyta fam., gen. et spec. indet. 3. D–F. Pteridophyta fam., gen. et spec. indet. 4. G–L. Pteridophyta fam., gen. et spec. indet. 5. Scale bars − 10 µm (A, B, D, E, G, H, J, K), 1 µm (C, F, I, L). 122 F. Grímsson et al. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 14. LM (A, D, G, J) and SEM (B, C, E, F, H, I, K, L) micrographs of dispersed fossil spores (A–F) and pollen (G–L). A–F. Pteridophyta fam., gen. et spec. indet. 6. G–L. Ginkgo sp. Scale bars − 10 µm (A, B, D, E, G, H, J, K), 1 µm (C, F, I, L). Sarmatian palynoflora from Lavanttal 123 Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Figure 15. LM (A, D, G, J) and SEM (B, C, E, F, H, I, K, L) micrographs of dispersed fossil pollen. A–C. Ginkgo sp. D–I. Ephedra sp. 1. J–L. Ephedra sp. 2. Scale bars − 10 µm (A, B, D, E, G, H, J, K), 1 µm (C, F, I, L). 124 F. Grímsson et al. (frontal view of longest diameter); equatorial diameter 41–46 µm wide in LM, 37–43 µm in SEM, polar axis 22–24 µm long in LM, 16–18 µm in SEM; polyplicate, seven plicae (0–7 intermediate plicae); exine 0.8–1.1 µm thick (LM); tectate; sculpturing psilate, margin of plicae slightly sinuous, plicae crested, closely spaced (SEM). Remarks. — Ephedra pollen have a solid pollen record dating back at least to the Early Cretaceous (Rydin et al., 2006b; Tekleva & Krassilov, 2009), and are frequently found in Cainozoic sediments of the Northern Hemisphere (Maher, 1964; Cookson, 1956; Zetter, 1988; Ferguson et al., 1998; Stuchlik et al., 2001; Denk et al., 2011). The Ephedra sp. 1 pollen type is comparable to pollen of many extant Eurasian Ephedra species, including E. gerardiana Wallich ex. C.A. Meyer, E. likiangensis Florin, E. przewalskii Stapf, and E. alata DC. (Reille, 1992; Fujiki et al., 2005). Fossil pollen grains similar to this type and the Ephedra sp. 2 type (see later) from the geological record have been assigned to the fossil genus Distachyapites or Ephedripites (Stuchlik et al., 2002). Ecological implications. — The genus Ephedra comprises 35–45 extant species with a wide distribution in temperate and warm arid regions of the Northern Hemisphere and in South America. There are c. 40 species in Eurasia, 12 in North America, and 13 in South America (Kramer & Green, 1990; Flora of North America Editorial Committee, 1993). In North America, Ephedra occurs in south-western USA and northern Mexico. There, the plants are growing on dry, more or less open sites that range in elevation from sea level up to c. 3000 m (cf. Maher, 1964). Ephedra shows a considerable ecological range, growing on well drained rocky to sandy substrate, dry slopes, flat sandy areas, alluvial fans (flood lands), and valley grassland, but it is also found on canyon walls and in ravines (Flora of North America Editorial Committee, 1993). In Asia, Ephedra occurs from the lowlands to high mountains of up to 4600 m. The Asian taxa also show a wide ecological range, growing mostly in sandy or rocky sites of flood lands and river valleys, on mountain slopes and cliffs but also on grasslands and in deserts (Flora of China Editorial Committee, 1999). Ephedra sp. 2 (Figure 15J–L) Description. — Pollen, monad, oblate, elliptic in polar view, star-like in equatorial view; equatorial diameter 40–43 µm wide in LM, 36–39 µm in SEM, polar axis 26–29 µm long in LM, 23–25 µm in SEM; inaperturate, polyplicate, seven plicae; exine 0.9–1.1 µm thick (LM); tectate; sculpturing psilate to foveolate, margins of plicae markedly sinuous, plicae rounded, widely spaced, area between plicae divided by foveolae into variably shaped units (SEM). Remarks. — The Ephedra sp. 2 pollen type is comparable to pollen of the extant E. intermedia Schrenk ex C.A. Meyer, E. distachya L., E. major Host., E. monostachya L., and E. helvetica (C.A. Meyer) Ascherson (Reille, 1992; Fujiki et al., 2005). Discussion Occurrence and identification of palynomorphs Spores are rare in the Lavanttal samples and each spore types are represented by less than 1% of the palynomorph spectrum. In a typical LM pollen count (300–450 grains), only one or two spore types may be encountered. This is also the case for the Ephedra grains, which could easily be missed in pollen counts. The only grains occurring with some abundance are those assigned to Ginkgo. Still, this pollen type also makes up less than 1% of the whole spectrum. For fossil spores of ferns, where the perispore is abraded, precise systematic assignment at generic or family level may be difficult, particularly when the sculpturing of the more robust exospore and the perispore are markedly different. This is the case for the most common spore type in the palynoflora described as Pteridophyta fam., gen. et spec. indet. 1 including psilate, kidney-shaped spores lacking the perispore. A similar general exospore morphology is known for more than ten families of ferns (Stuchlik et al., 2001). For other fern-like spores such as those of Lycopodium and Osmunda, in which the thin perispore is moulded on top of the more robust exospore sculpturing, determination is more secure even when the perispore is lacking. Of the 17 spore types presented here (Table I), ten are described for the first time from the Lavanttal Basin. The Ginkgo type pollen and the Ephedra sp. 1 pollen type are also reported for the first time. Even though there are many Ginkgo pollen grains in the samples, it is not surprising that Klaus (1984) did not report any since Ginkgo pollen is hardly identified by LM only and may be mistaken for various collapsed algal cysts. Ginkgo pollen may also be difficult to distinguish from pollen of Cycadaceae using LM only. Identification of Ginkgo pollen is more secure using SEM examination due to its very faint sculpturing that is otherwise obscured. Many fossil pollen grains that are described under the generic name Cycadopites could in fact be pollen from Ginkgo. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Sarmatian palynoflora from Lavanttal 125 Table I. Palynomorphs described in this study in comparison to Klaus (1984). This study Sphagnum sp. Lycopodium sp. 1 Lycopodium sp. 2 Selaginella sp. Dryopteris sp. Osmunda sp. Polypodiaceae gen. et spec. indet. 1 Polypodiaceae gen. et spec. indet. 2 Pteris sp. 1 Pteris sp. 2 Klaus (1984) Sphagnum sp. Osmunda sp. Verrucatosporites alienus Polypodium vulgare ? Polypodiacaeoisporites speciosus ? Pteris sp. ? Verrucingulatisporites sp. Polypodiaceaesporites haardtii Pteris sp. 3 Pteridophyta fam., gen. et spec. indet. 1 Pteridophyta fam., gen. et spec. indet. 2 Pteridophyta fam., gen. et spec. indet. 3 Pteridophyta fam., gen. et spec. indet. 4 Pteridophyta fam., gen. et spec. indet. 5 Pteridophyta fam., gen. et spec. indet. 6 Ginkgo sp. Ephedra sp. 1 Ephedra sp. 2 All of the extant taxa producing spores and pollen that are comparable to the fossils identified from the Lavanttal Basin are presently distributed in temperate to warm temperate regions. Some taxa have an extended distribution into cold-temperate environments and only a small minority extends into subtropical to tropical regions. In general, the taxa encountered so far are suggestive of a relatively warm and humid climate without dry winters. The taxa described here present only a small part of the palynoflora from the phosphoritic nodules of the Lavanttal Basin and detailed interpretations of the palaeo-vegetation and climate will be presented in the concluding part of this series of papers when all palynomorphs are properly documented. Acknowledgements The authors wish to thank the FWF (Austrian Science Foundation) for granting FG with a LiseMeitner-grant (project number M 1181-B17) that made this study possible. They are also grateful to Christa-Charlotte Hofmann, Hugh Rice, and Thomas Denk for reading over and improving the manuscript. The authors also thank the editor Else Marie Friis, David Batten, and another anonymous reviewer for good and constructive suggestions. Downloaded By: [Grímsson, Friðgeir] At: 08:59 18 June 2011 Ephedra distachya Note: Question marks indicate when we are uncertain if the LM based description and/or micrographs by Klaus (1984) correspond to our types. References Ambwani, K., Sahni, A., Kar, R. K. & Dutta, D. (2003). Oldest known non-marine diatoms (Aulacoseira) from the uppermost Cretaceous Deccan Intertrappean beds and Lameta Formation of India. Revue de Micropaléontologie, 46, 67–71. Bassir, S. H. (1964). Die Kohleflöze des Lavanttales. Leoben: University of Leoben, PhD Diss. Batten, D. J., Gray, D. J. & Harland, R. (1999). Palaeoenvironmental significance of a monospecific assemblage of dinoflagellate cysts from the Miocene Clarkia Beds, Idaho, USA. Palaeogeography, Palaeoclimatology, Palaeoecology, 153, 161–171. Batten, D. J. & Grenfell, H. R. (1996). Botryococcus. In J. Jansonius & D. C. McGregor (Eds), Palynology: principles and applications, Vol. 1 (pp. 205–214). Salt Lake City, UT: American Association of Stratigraphic Palynologists Foundation, Publishers Press. Bechtel, A., Reischenbacher, D., Sachsenhofer, R. F., Gratzer, R., Lücke, A. & Püttmann, W. (2007). Relations of petrographical and geochemical parameters in the Middle Miocene Lavanttal lignite (Austria). International Journal of Coal Geology, 70, 325–349. Beck-Mannagetta, P. (1952). Zur Geologie und Paläontologie des Tertiärs des unteren Lavanttales. Jahrbuch der Geologischen Bundesanstalt, 95, 1–102. Berger, W. (1953). Pflanzenreste aus den obermiozänen Ablagerungen von Wien-Hernals. Annalen Naturhistorisches Museum Wien, 59, 141–154. Berger, W. (1955). Jungtertiäre Pflanzenreste aus dem unteren Lavanttal in Ostkärnten. 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