Acta Palaeobotanica Early Oligocene plant diversity along the Upper Rhine Graben The fossil flora of Rauenberg Germany

Johanna  Eder

The macroflora of Rauenberg, Baden-Württemberg, Germany, is treated monographically. The plant-bearing sediments are marine, mainly well-bedded clay-to siltstones, the so-called Fischschiefer, which are part of the Bodenheim Formation. Based on nannoplankton they are dated to nannoplankton zone NP 23 (Rupelian, Lower Oligocene). The plant remains, mainly leaves and some fructifications, are preserved as compressions. The taxonomic assignment is based on gross morphology and cuticle characteristics. The flora yields marine algae and remains of the very diverse terrestrial flora. A total of 68 taxa, including three types of algae, one cycad, 12 conifers, and 49 dicots, among them 5 palms, are described. The following fossil species are described for the first time: Laurophyllum rauenbergense, Myrica obliquifolia, Distylium metzleri, ? Berchemia altorhenana, ? Ternstroemites maritiae, Trachelospermum kelleri, Oleinites altorhenana, O. rauen-bergensis, Dicotylophyllum badense, D. oechsleri, D. vesiculaeferens, D. ziegleri, ? Viscophyllum hendriksiae, and Cladites vesiculaeferens. Dicotylophyllum vesiculaeferens and Cladites vesiculaeferens bear peculiar, complex cuticular structures presumably representing salt-secreting glands. Both taxa are interpreted to derive from one plant species of yet uncertain systematic affinity. The flora bears a high proportion of broad-leaved, presumably evergreen taxa, whereas the diversity of modern Arcto-Tertiary taxa (sensu Kvaček 1994) is rather low. Most abundant are Platanus neptuni, Daphnogene cinnamomifolia, and Tetraclinis salicornioides. On the family level, Lauraceae (10 species) and Pinaceae (8) are most diverse, followed by Arecaceae (4–5), Cupressaceae, and Myricaceae (4 species each). Surprisingly, Fagaceae are documented solely by a single leaf of Eotrigonobalanus furcinervis f. haselbachensis, and the record of Pentaphyllaceae remains ambiguous (? Ternstroemites maritiae). Sloanea olmediaefolia is recorded for the first time from western parts of Europe. Remarkable is the presence of the rare cycad Ceratozamia floersheimensis. The following possible vegetation units are suggested: zonal broad-leaved sclerophyllous evergreen forests and an intrazonal coastal pine-laurel-palm association on near-coastal sandy soils, as well as gallery forests along streams. No records of swampy environments were recovered. The climate may be characterised as follows: Cfa climate in transition to Cwa (and Am or Af) climate sensu Köppen, mean annual temperature 19–24°C, mean annual precipitation 1300–1700 mm, mean temperature of the warm-est month 28–29°C, mean temperature of the coldest month 8–14°C, mean precipitation of the wettest month >230 mm, mean precipitation of the driest month 18–38 mm, wettest month between May and October, driest month between November and March. The warm period was the wetter one. The flora from Rauenberg most closely resembles that of Flörsheim (Kvaček 2004a) and shows relations to the Paratethys realm, for example the Tard Clay Formation. Relations to the floras from Saxony, Saxony-Anhalt, and North Bohemia, similar in age, are rather restricted: broad-leaved deciduous taxa are much less diverse, and the numerous presumably evergreen taxa and palms present in Rauenberg have not been recorded in the other regions, indicating a more complex vegetation differentiation than a simple north-south gradient. The high number of taxa of uncertain affinity at Rauenberg points to the need for further taxonomic studies of the flora of this time interval. Comparisons with European assemblages of the early Oligocene reveal that the vegetation diversity in Europe during this time interval is far from being well understood.

Acta Palaeobotanica 56(2): 329–440, 2016 DOI: 10.1515/acpa-2016-0011 Early Oligocene plant diversity along the Upper Rhine Graben: The fossil lora of Rauenberg, Germany JOHANNA KOVAR-EDER State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany; e-mail: johanna.eder@smns-bw.de Received 11 July 2016; accepted for publication 7 October 2016 ABSTRACT. The macrolora of Rauenberg, Baden-Württemberg, Germany, is treated monographically. The plant-bearing sediments are marine, mainly well-bedded clay- to siltstones, the so-called Fischschiefer, which are part of the Bodenheim Formation. Based on nannoplankton they are dated to nannoplankton zone NP 23 (Rupelian, Lower Oligocene). The plant remains, mainly leaves and some fructiications, are preserved as compressions. The taxonomic assignment is based on gross morphology and cuticle characteristics. The lora yields marine algae and remains of the very diverse terrestrial lora. A total of 68 taxa, including three types of algae, one cycad, 12 conifers, and 49 dicots, among them 5 palms, are described. The following fossil species are described for the irst time: Laurophyllum rauenbergense, Myrica obliquifolia, Distylium metzleri, ? Berchemia altorhenana, ? Ternstroemites maritiae, Trachelospermum kelleri, Oleinites altorhenana, O. rauenbergensis, Dicotylophyllum badense, D. oechsleri, D. vesiculaeferens, D. ziegleri, ? Viscophyllum hendriksiae, and Cladites vesiculaeferens. Dicotylophyllum vesiculaeferens and Cladites vesiculaeferens bear peculiar, complex cuticular structures presumably representing salt-secreting glands. Both taxa are interpreted to derive from one plant species of yet uncertain systematic afinity. The lora bears a high proportion of broad-leaved, presumably evergreen taxa, whereas the diversity of modern Arcto-Tertiary taxa (sensu Kvaček 1994) is rather low. Most abundant are Platanus neptuni, Daphnogene cinnamomifolia, and Tetraclinis salicornioides. On the family level, Lauraceae (10 species) and Pinaceae (8) are most diverse, followed by Arecaceae (4–5), Cupressaceae, and Myricaceae (4 species each). Surprisingly, Fagaceae are documented solely by a single leaf of Eotrigonobalanus furcinervis f. haselbachensis, and the record of Pentaphyllaceae remains ambiguous (? Ternstroemites maritiae). Sloanea olmediaefolia is recorded for the irst time from western parts of Europe. Remarkable is the presence of the rare cycad Ceratozamia loersheimensis. The following possible vegetation units are suggested: zonal broadleaved sclerophyllous evergreen forests and an intrazonal coastal pine-laurel-palm association on near-coastal sandy soils, as well as gallery forests along streams. No records of swampy environments were recovered. The climate may be characterised as follows: Cfa climate in transition to Cwa (and Am or Af) climate sensu Köppen, mean annual temperature 19–24°C, mean annual precipitation 1300–1700 mm, mean temperature of the warmest month 28–29°C, mean temperature of the coldest month 8–14°C, mean precipitation of the wettest month >230 mm, mean precipitation of the driest month 18–38 mm, wettest month between May and October, driest month between November and March. The warm period was the wetter one. The lora from Rauenberg most closely resembles that of Flörsheim (Kvaček 2004a) and shows relations to the Paratethys realm, for example the Tard Clay Formation. Relations to the loras from Saxony, Saxony-Anhalt, and North Bohemia, similar in age, are rather restricted: broad-leaved deciduous taxa are much less diverse, and the numerous presumably evergreen taxa and palms present in Rauenberg have not been recorded in the other regions, indicating a more complex vegetation differentiation than a simple north-south gradient. The high number of taxa of uncertain afinity at Rauenberg points to the need for further taxonomic studies of the lora of this time interval. Comparisons with European assemblages of the early Oligocene reveal that the vegetation diversity in Europe during this time interval is far from being well understood. KEYWORDS: leaves, cuticles, taxonomy, vegetation reconstruction, Rupelian, NP 23 Unangemeldet Heruntergeladen am | 10.04.17 15:37 330 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 CONTENTS Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geographical, geological setting and age . . . . . . Materials and methods . . . . . . . . . . . . . . . . . . . . Systematic part. . . . . . . . . . . . . . . . . . . . . . . . . . . Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microthyriaceae . . . . . . . . . . . . . . . . . . . . Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phaeophyceae . . . . . . . . . . . . . . . . . . . . . . Phaeophyceae vel Rhodophyceae . . . . . . . Ferns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizaeales . . . . . . . . . . . . . . . . . . . . . . . . Schizaeaceae . . . . . . . . . . . . . . . . . . . . Gymnosperms . . . . . . . . . . . . . . . . . . . . . . . . . Cycadales . . . . . . . . . . . . . . . . . . . . . . . . . Zamiaceae . . . . . . . . . . . . . . . . . . . . . . Araucariales/Cupressales . . . . . . . . . . . . Doliostrobaceae . . . . . . . . . . . . . . . . . . Cupressales . . . . . . . . . . . . . . . . . . . . . . . . Cupressaceae . . . . . . . . . . . . . . . . . . . . Pinales . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pinaceae . . . . . . . . . . . . . . . . . . . . . . . . Angiosperms . . . . . . . . . . . . . . . . . . . . . . . . . . Laurales . . . . . . . . . . . . . . . . . . . . . . . . . . . Lauraceae . . . . . . . . . . . . . . . . . . . . . . . Arecales . . . . . . . . . . . . . . . . . . . . . . . . . . . Arecaceae . . . . . . . . . . . . . . . . . . . . . . . Proteales. . . . . . . . . . . . . . . . . . . . . . . . . . . Platanaceae . . . . . . . . . . . . . . . . . . . . . Buxales. . . . . . . . . . . . . . . . . . . . . . . . . . . . ? Buxaceae . . . . . . . . . . . . . . . . . . . . . . Fagales . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fagaceae . . . . . . . . . . . . . . . . . . . . . . . . Betulaceae . . . . . . . . . . . . . . . . . . . . . . 330 331 333 333 333 333 333 333 333 334 334 334 334 334 334 335 335 335 335 336 336 338 338 338 343 343 345 345 346 346 346 346 357 Betulaceae vel Ulmaceae . . . . . . . . . . Myricaceae . . . . . . . . . . . . . . . . . . . . . . Juglandaceae . . . . . . . . . . . . . . . . . . . . ? Rosales. . . . . . . . . . . . . . . . . . . . . . . . . . . ? Rhamnaceae . . . . . . . . . . . . . . . . . . . ? Fabales . . . . . . . . . . . . . . . . . . . . . . . . . . ? Fabaceae . . . . . . . . . . . . . . . . . . . . . . Oxidales . . . . . . . . . . . . . . . . . . . . . . . . . . . Elaeocarpaceae. . . . . . . . . . . . . . . . . . . Malphigiales. . . . . . . . . . . . . . . . . . . . . . . . Salicaceae . . . . . . . . . . . . . . . . . . . . . . . Malvales . . . . . . . . . . . . . . . . . . . . . . . . . . . Malvaceae . . . . . . . . . . . . . . . . . . . . . . Saxifragales . . . . . . . . . . . . . . . . . . . . . . . . Hamamelidaceae . . . . . . . . . . . . . . . . . ? Santalales . . . . . . . . . . . . . . . . . . . . . . . . ? Loranthaceae . . . . . . . . . . . . . . . . . . . Cornales . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrangeaceae . . . . . . . . . . . . . . . . . . Ericales. . . . . . . . . . . . . . . . . . . . . . . . . . . . ? Pentaphyllaceae . . . . . . . . . . . . . . . . Symplocaceae . . . . . . . . . . . . . . . . . . . . Gentianales . . . . . . . . . . . . . . . . . . . . . . . . Apocynaceae . . . . . . . . . . . . . . . . . . . . . Lamiales. . . . . . . . . . . . . . . . . . . . . . . . . . . Oleaceae . . . . . . . . . . . . . . . . . . . . . . . . Fam. indet. . . . . . . . . . . . . . . . . . . . . . . . . Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Flora of Rauenberg in the European context References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347 347 350 351 351 352 352 352 352 354 354 354 354 354 354 355 355 356 356 357 357 357 358 358 360 360 362 367 370 375 384 390 436 INTRODUCTION During the early Oligocene, Earth’s ecosystems adapted to cooling climatic conditions following the preceding Late Eocene thermal maximum (Zachos et al. 2008, Roth-Nebelsick et al. 2014). In Europe, diverse loras of this period are known from the southeastern margin of the Paratethys Sea (Hungary, Tard Clay Formation; Kvaček & Hably 1998, Hably & Erdei 2015) and the northern margin of former Europe, that is, North Bohemia (Bechlejovice (Kvaček & Walther 2004), Kundratice (Kvaček & Walther 1998), Markvartice (Bůžek et al. 1976, Kvaček & Walther 2001), Suletice-Berand (Kvaček & Walther 1995)) and Saxony (Hammerunterwiesenthal (Walther 1998), Haselbach Serie (Mai & Walther 1978), Kleinsaubernitz (Walther 1999), and Seifhennersdorf (Walther & Kvaček 2007)). In the Upper Rhine valley and the Mainz Basin, early Oligocene marine sediments are known for their rich fossil record. From the Mainz Basin, Kvaček (2004a) revised the plant record from marine deposits of Flörsheim, and Pross et al. (1998) studied palynomorphs (Bodenheim drill core), both NP 23, Rupelian in age. In the region around the town of Rauenberg near Wiesloch, these clay- to siltstones were mined in several clay pits. In the respective museum collections, such material is usually held under the locality name Frauenweiler, which is located in the northwestern Unangemeldet Heruntergeladen am | 10.04.17 15:37 331 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 part of Rauenberg. This study follows the proposal of Maxwell et al. (2016) and simply summarises all material from different clay pits under Rauenberg. While ish, e.g., Micklich & Parin (1996), dinolagellate cysts, calcareous nannoplankton and foraminifera (Grimm et al. 2002), and birds, e.g., Mayr (2004), Mayr & Micklich (2010), have been studied in detail, this is not the case for other organism groups such as molluscs, insects, reptiles, and plants. A short note on the macro-lora was published by Winterscheid & Gregor (2008). In the synthesis of the Rauenberg fossil lagerstätte, Maxwell et al. (2016) provide a comprehensive summary of all organismic groups present at Rauenberg, including a preliminary list of fossil plant taxa and the ecological implications of this lora. Since most of the plant taxa are entire-margined, a taxonomic assessment requires the application of cuticular analysis. The results of those taxonomic studies are provided in the present paper. Due to the high organismic diversity, the fossil lagerstätte of Rauenberg offers a unique insight into an early Oligocene ecosystem (Maxwell et al. 2016). The State Museum of Natural History Stuttgart (SMNS) houses a large fossil collection from Rauenberg including plants, invertebrates (insects, molluscs, crustaceans) and vertebrates (ish, reptiles, birds). These are available thanks mainly to the private collectors Rudolf Metzler and Harald and Annette Oechsler, who have conducted ieldwork over many years. Additional material derives from museum ield campaigns. The collection of fossil plants comprises over 650 slabs. In the course of this study, the respective collection in the Naturmuseum Augsburg (NMA) was also examined, which was the basis for the publication by Winterscheid & Gregor (2008), as well as the collection at the Staatliches Museum für Naturkunde Karlsruhe (SMNK). Comparative studies of the cuticle slides from the lora of Flörsheim (Kvaček 2004a) were limited because these slides are almost completely dried. GEOGRAPHICAL AND GEOLOGICAL SETTING AND AGE The town of Rauenberg is located 16 km south of Heidelberg, State of Baden-Württemberg, Germany, along the Upper Rhine valley. The clay pits around Rauenberg are situated in the Upper Rhine Graben (Fig. 1), a region of intensive tectonics during the Palaeogene and Neogene and still active today. The sediments exposed in the clay pits stem from the second Rupelian transgression (Hardenbol et al. 1998). They are assigned to the marine to brackish Froidefontaine Subgroup, Bodenheim Formation, which extends widely in the Upper Rhine Graben and its subbasins (Grimm 2005, Grimm et al. 2011). In the Rauenberg clay pits the Wallau and Hochberg Members of the Bodenheim Formation are developed. Lithologically the sediments are partly well-bedded or laminated clay- to siltstones. Rhythmic sedimentological events and changing carbonate content resulted in lamination, among other processes. Frequent mass occurrences of calcareous nannoplankton produce laminae of calcareous mud or calcareous sand layers. Based on calcareous nannoplankton, the Hochberg Member is assigned to NP 23. Rupelian age is supported by dinolagellate cysts and foraminifera (Grimm et al. 2002). The Alzey Formation, which is characterised by coarser sediments, represents the fully marine coastal equivalent of the Bodenheim Formation and comprises sands and gravels (Grimm 2005). Barite concretions in these sandy sediments of the Alzey Formation bear, among other fossils, abundant remains of pine cones and wood (Mallison 2002). For detailed geology and stratigraphy see Maxwell et al. (2016). MATERIALS AND METHODS The fossil lora of Rauenberg yields mainly leaves and more rarely fructiications, as well as marine algae. The plant material is preserved mainly as carbonised compressions and is highly scattered, as is characteristic of marine sediments. Private collectors preferred to collect complete specimens rather than very fragmented ones. Such material has been collected only since 2007, when the author contacted the collectors. Little stratigraphic information is available for the collection material from private collectors. Accordingly, potential changes in composition in the geological section cannot be traced. This information has been rigorously documented only since the SMNK began ield campaigns. The plant material was studied gross morphologically and, if possible, by cuticular analysis. Unfortunately, much of the material collected prior to 2007 either is coated by varnish or has been transferred to an artiicial resin, both hampering cuticle studies. Unangemeldet Heruntergeladen am | 10.04.17 15:37 332 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 A B North Sea Rhenish Massif Germany W Fl Rhine Graben Ra North European Platform Be,S,K Bohemian Massif Frankfurt Mannheim Nuremberg Rauenberg L Bu,E-K Massif Central Stuttgart Munich PRIABON. BARTON. LUTETIAN Upper Hydrobia Beds Wiesbaden, Rüssingen Fm. Upper Cerithium Beds Middle Cerithium Beds Molasse alsacienne Lower Cerithium Beds Freshwater Beds Freshwater Beds Serie grise Serie grise Alzey, Bodenheim, Alzey, Bodenheim, Stadecken Fm. Stadecken Fm. Upper Salt Fm. Haustein Pechelbronn Beds Middle Salt Fm. Plattiger Steinmergel Lower Salt Fm. Melanienkalke Lower Salt Fm. Melanienkalke Lymneenmergel Lymneenmergel Planorbenkalke Planorbenkalke 27 28 29 30 31 Chattian Ma 26 NP25 D15 c MP26 b MP25 a MP24 NP24 Rupelian RUPELIAN D Upper Rhine Graben North Oligocene UPPER CHATTIAN UPPER LOWER EOCENE OLIGOCENE AQUITAN. Upper Rhine Graben South Upper Rhine Graben conglomerates Series Stage Upper Rhine Graben conglomerates C C al c N are an o no us fo ss D in ils o Zo cy s ne t Eu M rop am ea M m n N a Zo -M ls ne P s Western Tethys b MP23 NP23 D14 32 a MP22 D13 D12 c MP20 * NP22 33 NP21 MP21 * Hochberg Member (Fischschiefer) Rauenberg Fig. 1. Geography and stratigraphy. A – Early Oligocene (Rupelian) palaeogeography of Europe, showing important Oligocene localities discussed in the text. Map modiied from Meulenkamp et al. (2000) based on Spiegel et al. (2007). B – Location of Rauenberg, Baden-Württemberg, Germany. C – Lithostratigraphy of the Upper Rhine Graben, modiied after Rasser et al. (2008). D – Estimated biostratigraphic age of the Hochberg Member (Fischschiefer). Fl – Flörsheim, Mainz Basin; Ra – Rauenberg; S – Seifhennersdorf; W – Weisselster Basin, Germany; Be – Bechlejovice; K – Kundratice, Czech Republic; Bu, E-K –Budapest and Eger-Kiseged, Hungary; L – Linz, Austria. Figure 1 after Maxwell et al. (2016) slightly modiied Nevertheless, cuticles were prepared from about 300 specimens, of which >200 attempts were successful. To prepare the cuticles, the samples were treated with 5–10% hydrochloric acid (HCl) followed by 10–20% hydroluoric acid (HF) to dissolve the sediment. The leaf fragments were then macerated with Schulze´s reagent (HNO3+KClO3), followed by treatment in 5% potassium hydroxide solution (KOH). The cuticles were stained with safranin and embedded in glycerol. The cover glasses were sealed with nail polish. The cuticles of many specimens are heavily infected by fungi, often masking the stomata. For cuticle descriptions, representative values were obtained by measuring at least three leaves per taxon wherever possible. Ten measurements of every cell type (stomata, non-modiied epidermal cells, trichome bases per leaf) were taken wherever possible. Averages refer to values per leaf. Cuticles were studied under a Leitz interference contrast microscope. SEM studies were performed on a Zeiss EVO LS 15. All cuticle slides are stored in the SMNS. Acronyms in this study: NMA – Naturmuseum Augsburg, SMNK – Staatliches Museum für Naturkunde Karlsruhe, SMNS – State Museum of Natural History Stuttgart. Abbreviations in descriptions: l × w for length × width, l/w ratio for ratio length/width. Unangemeldet Heruntergeladen am | 10.04.17 15:37 333 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Microthyriaceae Sacc. tiny aerocysts. In marine deposits of different ages, Cystoseirites-type algae are sometimes frequent, for example in the surroundings of Linz, Austria, earliest Miocene (Kovar 1982), and Sarmatian deposits of Romania (Givulescu 1975, Paraschiv 2004). From Rauenberg this is the only remain; from Flörsheim (Mainz Basin) no aerocyst-bearing algae have been reported. Microthyriaceae gen. et sp. indet. Phaeophyceae vel Rhodophyceae SYSTEMATIC PART Systematics follows the Open Tree of Life (Hinchliff et al. 2015). FUNGI Pl. 8, igs 1–3 Laminarites Sternberg M a t e r i a l. SMNS P 1952/473. D e s c r i p t i o n. Cylindrical spores with rounded poles and concentrically growing fruiting bodies in different stages. R e m a r k s. Similar spores and fruiting bodies are met frequently on fossil leaf material, e.g., Kovar (1982). In Rauenberg they occur sporadically, while hyphae of fungi are much more common. From Upper Oligocene sediments in Greenland, Worobiec and Worobiec (2013) treated similar structures taxonomically. The specimens from Rauenberg are not assigned more closely because diagnositic features are poor. ALGAE Phaeophyceae Cystoseirites Sternberg Cystoseirites communis Unger Pl. 1, ig. 1 1847 Cystoseirites communis, Unger, p. 125, pl. 38, igs 1, 2. M a t e r i a l. SMNS P 1953/111. D e s c r i p t i o n. Filamentous, multiply branched thallus, ilaments up to 1 mm wide, bearing elongated aerocysts of up to 4 mm length and about 1 mm width. R e m a r k s. The aerocysts account for the assignment to the Phaeophyceae. Cystoseirites communis was irst described from Radoboj, Croatia (middle Miocene, Sarmatian). These algae differ from Cystoseirites altoaustriacus Kovar by having elongated, rather Laminarites latus Engelhardt Pl. 1, ig. 4 2004a Laminarites latus Engelhardt; Kvaček, p. 3, pl. 1, igs 9, 10. M a t e r i a l. SMNS P 1952/97 (together with Platanus neptuni), 188, 197, 199, 207, 213+214, 242, 295, 347. D e s c r i p t i o n. Fragments of thalli, bandlike, straight to gently bent, parallel-sided, densely spaced strands sometimes faintly visible, up to 160 mm long, up to 17 mm wide, unbranched. R e m a r k s. These thalli differ from the belowdescribed ones by their larger width and the absence of branchings. In specimen SMNS P 1952/242 the margins are darker, possibly indicating that the margin was slightly thicker. Thallites Walton Thallites multiidus (Brongniart) Kvaček Pl. 1, igs 2, 3 2004a Thallites multiidus (Brongniart) comb. nov.; Kvaček, p. 4, pl. 1, ig. 1. M a t e r i a l. SMNS P 1952/1, 39, 78, 126, 131, 138, 148, 323; SMNS P 1953/104, 121, 122. D e s c r i p t i o n. Thalli (multiply) dichotomously forked, sometimes forming dense mats, branches up to 3 mm wide. R e m a r k s. Remains of this type are quite common. From Flörsheim one small specimen has been described bearing a close resemblance based on the few available characters. Unangemeldet Heruntergeladen am | 10.04.17 15:37 334 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 FERNS Schizaeales termination ambiguous, margin dentate or crenulate, somewhat rolled into the sediment. Lygodium Sw. R e m a r k s. This pinna fragment differs from the above-described ones by the vein pattern in which the midvein is straight and the lateral veins arise under wider angles. Lygodium kaulfussii Heer emend. Gardner et Ettingshausen GYMNOSPERMS Schizaeaceae Kaulf. Pl. 1, igs 5–7 1861 Lygodium kaulfussi, Heer 1861, p. 409, pl. 9, ig 1. 2004a Lygodium kaulfusii Heer emend. Gardner et Ettingshausen; Kvaček, p. 4, pl. 1, ig. 11. Cycadales Zamiaceae Reichenbach Ceratozamia A. Brongniart M a t e r i a l. SMNS P 1952/55 (pinna), 322, 332 (isolated lobes). Ceratozamia loersheimensis (Engelhardt) Kvaček D e s c r i p t i o n. A single palmate pinna and two fragments; palmate pinna length × width (hereafter l × w) = 12 × 15 mm, base cuneate, pinna dissected into four lobes, three of them rather complete, lobes 2 mm wide, neither margin nor apices preserved; veins ascending steeply and forking, midvein vague; two isolated pinnae (i.e. lobes) both lacking base and apex, l × w = 21 × 5 mm (SMNS P 1952/322) and l × w = 19 × 4 mm (SMNS P 1952/332), narrow with parallel, inely crenulate/serrate margins, main vein slender, smoothly wavy, higher-order veins ascending steeply from the main vein, then turning towards the margin, course wavy, forking at least once, usually twice or even three times, ending in the marginal tooth apices. Pl. 1, ig. 8; Pl. 8, igs 4–7 R e m a r k s. Compared to the records from Flörsheim (Kvaček 2004a) and Geiseltal (Eocene, Barthel 1976), this is a remarkably tiny palmate pinna. Its margin is not preserved, thus the termination of the veins remains unclariied and the pinna lacks unambiguous main veins running into the lobes. The latter fact may, however, be an artefact caused by poor preservation, whereas the isolated lobes are better preserved. Filicopsida sp. M a t e r i a l. SMNS P 1952/69. D e s c r i p t i o n. Fragment of pinna, l × w = 9 (incomplete) × 3 mm, with distinct midvein, further veins arising from it at about 45° angle or more, once or twice forking, marginal 1911 Iris loersheimensis Engelhardt, p. 319, pl. 37, ig. 27. 2002 Ceratozamia loersheimensis (Engelhardt) comb. nov.; Kvaček, p. 305, igs 1–21. 2014 Ceratozamia loersheimensis (Engelhardt) Kvaček; Kvaček, p. 234, pl. 1, igs 6–7, pl. 2 ig. 10. M a t e r i a l. SMNS P 1952/456. D e s c r i p t i o n. Fragment of a lealet, texture coriaceous, l × w = 50 (incomplete) × 8 mm, narrow, parallel-margined, gradually tapering towards one end; venation parallel. M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle thick, bearing cells of two types: (1) elongated ones, anticlines straight to somewhat bent, length very variable up to 150 µm, width 10–18 µm, cross-walls mainly distinctly oblique, curved or S-shaped, (2) short rows (100 to at least 150 µm long) of more strongly cutinised short cells, length variable up to 50 µm, width 9–13 µm, epicuticular striation to folding paralleling the cell rows; stomatal complexes conined to abaxial cuticle, mainly longitudinally oriented, mono- to di- (or tri-) cyclocytic, subsidiary cells up to 6, very variable in shape, e.g., l × w = 93 × 71 µm, 114 × 74 µm), stomata somewhat recessed, remains of guard cells occasionally still present, e.g., l × w = 40 × 34 µm, 34 × 29 µm. R e m a r k s. This is the fragment of a narrow cycad lealet lacking base and apex, although it tapers at one end. It was detected among the material accessed since 2007, when collection of strongly fragmented material was encouraged Unangemeldet Heruntergeladen am | 10.04.17 15:37 335 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 by the author. Among cycads, short, thickly cutinised cells arranged in rows are characteristic of the genus Ceratozamia (Kvaček 2002). Diagnostically relevant features to distinguish C. loersheimensis and C. hofmannii are few. Of these, the presence of venation forkings remains open in the specimen from Rauenberg because the fragment is rather small. The lealet attachment also remains unclear because the base is missing. Based on cuticular features, C. hofmannii differs from C. loersheimensis by having distinctly smaller, short cells (Kvaček 2004b). In this respect, the here-described specimen clearly matches C. loersheimensis. Apart from Rauenberg and the type locality Flörsheim, C. loersheimensis is known from the Oligocene of Hungary (Budapest-Nagybátony, Tard Clay) and Slovenia (Trbovlje) (Kvaček 2002, 2014). A modern relationship exists to C. microstrobila Vovides & J.D. Rees, C. morettii Vázquez-Torres & Vovides, C. latifolia Miquel, and C. delucana Vázquez-Torres, Moretti & Carvajal-Hernández (Kvaček 2014). Araucales/Cupressales Doliostrobaceae Kvaček Doliostobus Marion Doliostobus taxiformis (Sternberg) Kvaček ? taxiformis Mai & Walther Pl. 1, ig. 9 1833 Cystoseirites taxiformis, Sternberg, fasc. 5–6, p. 35, pl. 18, igs 1–3. 1971a Doliostrobus taxiformis (Sternberg) comb. nov.; Kvaček, p.118, pl. 31, ig. 19, pl. 32, igs 1–5. 2008 Doliostrobus taxiformis (Sternberg 1833) Kvaček 1971; Winterscheid et Gregor, p. 4. M a t e r i a l. NMA 2006-104/1497. D e s c r i p t i o n. Terminate foliated shoot, branched twice near base, 110 mm long, tapering towards apex; leaves helically arranged, basally decurrent along axis, apical part free, awl-shaped. M i c r o m o r p h o l o g y. Large, straight-walled non-modiied epidermal cells of possibly stomabearing zones, presence of stomata ambiguous. R e m a r k s. For this specimen, cuticle preparation was not successful. Gross morphologically it is more similar to Doliostrobus taxiformis than to Chamaecyparites hardtii (Göppert) Endlicher in which the needles usually appear to be less coriaceous and less regularly arranged (compare e.g., Kvaček 1971a, pl. 31, ig. 17, Kunzmann 1999, pl. 24, igs 1–4). The assignment to the variety taxiformis, which is characterised by larger stomata than the variety sternbergii (Mai & Walther 1985), remains somewhat ambiguous. Winterscheid and Gregor (2008) listed D. taxiformis from Rauenberg without a igure. Although the collection ile number given in that publication does not correspond to the number on the specimen itself, I assume that those authors referred to this specimen for two reasons: (1) it is the only specimen in the respective collection in the Naturmuseum Augsburg that may be assigned to Doliostrobus; and (2) in their publication, numerous collection ile numbers do not correspond to those on the specimens themselves. Doliostrobus taxiformis is one of the ancient taxa in this assemblage. Cupressales Cupressaceae Sequoia Endl. Sequoia abietina (Brongniart in Cuvier) Knobloch Pl. 1, ig. 10; Pl. 8, ig. 8 1964 Sequoia abietina (Brongn. in Cuv.) Knobl., Knobloch, p. 601. M a t e r i a l. NMA 2016-3/2179. D e s c r i p t i o n. Foliated shoot 70 mm long, needles densely spaced, helically arranged but oriented in one plane along the twig, needles up to 11 mm long and 1.1–1.4 mm wide, entiremargined, needle width tapering towards base, twisted and somewhat decurrent along the twig, lacking a distinct petiole, apex acute acuminate. M i c r o m o r p h o l o g y. Non-modiied epidermal cells elongate, mainly rather rectangular to oblique, arranged in rows paralleling the needle length, stomata conined to stoma bands, mainly oriented paralleling the rows of non-modiied epidermal cells, stomata cyclocyclic to amphicyclic, subsidiaries more strongly cutinised than non-modiied cells, well staining, polar cells smaller than lateral ones, stomatal aperture narrow oblong, 19–29 (average 26) µm long. Unangemeldet Heruntergeladen am | 10.04.17 15:37 336 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 R e m a r k s. Gross morphologically the needles of Sequoia are usually wider than those of Taxodium. The prevailing orientation of the stomata parallel to the needle length unequivocally conirms this assignment. This is the only specimen documenting this species at Rauenberg. Taxodium Rich. Taxodium sp. Pl. 1, igs 11–13 M a t e r i a l. SMNS P 1952/88, SMNS P 1953/103. SMNK 7710. D e s c r i p t i o n. Detached shoots with longest needles in the middle part, needles helically inserted but spread in two opposite ranks, directly attached to the axis (sessile, no petiole), l × w up to 9 × 1 mm. R e m a r k s. The assignment to Taxodium is based on shoot shape, needle arrangement and needle width. Abietoid shoots of Sequoia are more robust and bear thicker needles. Twigs with cupressoid foliage reminiscent of Sequoia or Glyptostrobus are absent in Rauenberg. A further, poorly preserved, rather thick twig fragment shows cupressoid foliage and is assigned to the Taxodioideae. Records of the Taxodioideae are remarkably scarce in Rauenberg and are restricted to shoots. usually less than 10 mm; specimen SMNS P 1953/109 a twig fragment consisting of four whorls about 15 mm long and branching buds. M i c r o m o r p h o l o g y. Cuticle thick, anticlines of non-modiied epidermal cells distinct, undulate with knobs, forming quadrangular cells in non-stomatal areas and more irregular polygonal cells in stomatal areas, often one or two papillae (strongly cutinised regions) on the surface of the non-modiied epidermal cells, very variable in size, quadrangular cells up to 50 µm in length, polygonal cells 25–40 µm in diameter, stomatal complexes partly incompletely amphicyclocytic, adjacent stomata sometimes sharing subsidiary cells, aperture bordered by a thick, rather quadrangular Florin-ring, aperture diameter 6–12 µm. R e m a r k s. The true organisation of the leaf arrangement, namely four leaves in one whorl (not decussate) and marginally adnate, was recognised by Kovar (1982), although she hesitated to assign these remains to Tetraclinis. The twig SMNS P 1953/109, reminiscent of T. brachyodon (Brongniart) Mai & Walther (pl. 1, ig. 15), is assigned to T. salicornioides because different positions on the living plant may have been relected in morphological variability. Pinales Pinaceae Lindl. Tetraclinis Mast. Tetraclinis salicornioides (Unger) Kvaček Pinus L. Pl. 1, igs 14, 15; Pl. 8, ig. 9 Pinus (Strobus) sp. 1 1841 Thuytes salicornioides Unger, p. 11 pl. 2 igs 1–4. 1989 Tetraclinis salicornioides (Unger) comb. nov. ; Kvaček, p. 48–51, pl. 1 ig. 11, pl. 2 igs 2–14, pl. 3, igs 3, 4. 2008 Tetraclinis salicornioides (Unger 1841) Kvaček 1989; Winterscheid & Gregor, pl. 1 ig. 2. Pl. 1, ig. 16 M a t e r i a l. SMNS P 1952/20, 21, 70, 118, 144, 187, 194, 198, 226, 262, 294, 300, 313, 324, 326, 417, 420, 421, 422, 428, 457; SMNS P 1953/109. NMA 2006-118/1497. D e s c r i p t i o n. Fragments of twigs, most of them consisting only of one whorl of leaves fused in four, in some specimens several whorls still attached to each other, remains of branched twigs exceptional; whorl length M a t e r i a l. SMNS P 1952/292. ? SMNK 7747. D e s c r i p t i o n. Five-needled fascicle, needles 33 mm long, complete, slender, with a central keel, apex acute. R e m a r k s. Though the sheath is hardly preserved, this remain corresponds very well with the single fascicle of P. (Strobus) sp. described from Flörsheim by Kvaček (2004a). Pinus (Strobus) sp. 2 Pl. 1, ig. 17 M a t e r i a l. SMNS P 1952/89+90, ? 91, 248. Unangemeldet Heruntergeladen am | 10.04.17 15:37 337 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 D e s c r i p t i o n. Five-needled fascicles, needles 72 mm (SMNS P 1952/89+90) to 89 mm (SMNS P 1952/248) long, slender, with a central keel, apex acute. R e m a r k s. These fascicles differ from the foregoing species by having distinctly longer needles. In specimen SMNS P 1952/248, four needles are visible, and in specimen SMNS P 1952/91 only three; the others probably are covered by sediment. Pinus sp. 6 Pl. 1, ig. 20 2008 Pinus sp. – dreinadelige Belaubung; Winterscheid & Gregor, p. 4. M a t e r i a l. NMA 2006-111/1497. D e s c r i p t i o n. Single, almost complete, threeneedled fascicle, full needle length 66 mm. R e m a r k s. This is the only record of a threeneedled pine species. Pinus sp. 3 Pinus engelhardtii Menzel Pl. 1, ig. 18 Pl. 3, ig. 10 M a t e r i a l. SMNS P 1952/92, 460. NMA 2006103/1497. D e s c r i p t i o n. Two-needled fascicles, up to 115 mm long, up to 1 mm wide, rigid, apex ?, sheath up to ? 5 mm long. R e m a r k s. The needles appear to be rather delicate but stiff. Pinus sp. 4 M a t e r i a l. SMNK 7706. D e s c r i p t i o n. Slender elliptic cone, l × w = 83 (? 85–90) × about 40 mm, cone scales curved strongly outwards, apophyses narrow-rhomboidal, with short umbo and mucro. R e m a r k s. This cone differs markedly from the below-described ones by its distinctly recurved cone scales with narrow apophyses. Pl. 1, ig. 19 Pinus (Pinus) cf. thomasiana (Goeppert) Reichenbach M a t e r i a l. SMNS P 1952/143. D e s c r i p t i o n. Pair of needles 155 mm long (incomplete), 1.5 mm wide, base and apex incomplete, needles gently bent. R e m a r k s. This species differs from the foregoing one by its broader and probably longer needles. Pinus sp. 5 2008 Pinus sp. – zweinadelige Belaubung; Winterscheid & Gregor, p. 4, pl. 1, ig. 1. M a t e r i a l. NMA 2006-101/1497, 2006-102/ 1497. D e s c r i p t i o n. Two-needled fascicles, needle length 72 mm (complete)–80 mm or somewhat more. R e m a r k s. This species differs from Pinus spp. 3 and 4 by having shorter needles. Specimen NMA 2006-101/1497 has been igured by Winterscheid and Gregor (2008, pl. 1 ig. 1) under an incorrect inventory number: 2006184/1497. Pl. 3, igs 11, 12 Pinus nodosa Ludwig; Mallison, p. 19, pl. 3 igs 6–8. 2004a ? Pinus (Pinus) cf. thomasiana (Goeppert) Reichenbach; Kvaček, p. 6, pl. 3 igs 10, 11, 13. 2002 M a t e r i a l. SMNS P 1952/? 48, 176; SMNS P 1953/106, 110. D e s c r i p t i o n. Broad conical cones, l × w = 40–60 × 28–37 mm, petiole adherent, cone scales thick, apophyses rather lat, rhomboidal, wider than high, with medial keel, (ex)centromucronate; towards cone apex, keel and umbo shifting towards top of the apophyses. R e m a r k s. These specimens have been embedded by transfer technique in artiicial resin. It is likely that specimens SMNS P 1952/176 and SMNS P 1953/110 represent part and counterpart shared between the private collectors and later placed in the collection of the SMNS. In all specimens the apophyses are rather lat, as is characteristic for Pinus hampeana, which is known only from the Miocene. The Palaeogene species P. thomasiana (Goeppert) Reichenbach bears a resemblance, but the apophyses are Unangemeldet Heruntergeladen am | 10.04.17 15:37 338 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 domed, whereas they are relatively lat in our specimens. Whether this state is an artefact due to diagenetic processes remains unclear. From baryth concretions of the Alzey Formation, Pinus nodosa Ludwig (Mallison 2002) resembles the here-described pine cones, although details of the apophyses and umbo are less distinctly preserved in these specimens. Concluding from the assignment, the pine cones from Flörsheim were also clearly dificult to determine (Kvaček 2004a). P. thomasiana is assigned to section Sylvestres Van der Burgh, Series Nigrae Novák, showing close afinities especially to modern P. nigra Arnold and P. heldreichii Christ (Mai 1986). ANGIOSPERMS Laurales Lauraceae Juss. asymmetric, stomata sunken, stoma l × w = 15–19 (average 17) × 12–18 (average 14) µm, stomatal aperture narrow, slit-like, ledges somewhat thickened, pore short, mesophyllous oil cells abundant, variable in size (7–66 µm). R e m a r k s. The complete size of the heredescribed specimen can be only roughly estimated. Trichome bases have not been observed. The stomata may resemble Actinodaphne pseudogermari Walther & Kvaček and Laurophyllum acutimontanum Mai. In L. meuselii the stomata are somewhat diffusely overlapped by the delicate surface of the adjacent cells, while in A. pseudogermari the overlap is sharply delimited. L. acutimontanum differs both from A. pseudogermari and L. meuselii by its mostly stronger overlap of the stomata by domed cells leaving a rather narrow, irregularly slit-like opening above the stoma. The margins of the central opening appear somewhat thickened, unlike L. meuselii. Previously, A. pseudogermari was known only from the Upper Eocene. Actinodaphne Nees Daphnogene Unger Actinodaphne pseudogermari Walther in Mai & Walther 1985 Daphnogene cinnamomifolia (Brongniart) Unger Pl. 2, ig. 26; Pl. 9, igs 1–3 1985 Actinodaphne pseudogermari Walther nov. spec.; Mai & Walther, p. 33, pl. 22, igs 1–4, Bild 1/12–15, Bild 13/3–4. 2002 Actinodaphne pseudogermari Walther in Mai & Walther, 1985; Kunzmann & Walther, p. 266, igs 4a, g, 5d, e. M a t e r i a l. SMNS P 1952/17. D e s c r i p t i o n. Fragment of a leaf base with a short and swollen petiole, midvein prominent, petiole and midvein with distinct wrinkles parallel to the leaf axis, laminar shape ambiguous, base angle acute, base straight but convex directly above the petiole and somewhat asymmetric; l × w = 65 (? 120) × 25 (? 26–27) mm, entire-margined, secondary veins arising at an angle of up to 45°. M i c r o m o r p h o l o g y. Adaxial and abaxial cuticles thick, glabrous; adaxial cuticle: anticlines very thick, curved to somewhat wavy, non-modiied cells 17–26 (average 22) µm across; abaxial cuticle: anticlines distinct but less thick than adaxially, straight to curved, size of non-modiied cells 17–23 (average 21) µm, stomatal complexes brachyparacytic, Pl. 2 igs 1–6; Pl. 9 igs 4–9 Daphnogene cinnamomifolia (Brongniart) Unger; Mallison p. 39, pl. 9, igs 3, 4. 2004a Daphnogene cinnamomifolia (Brongniart) Unger; Kvaček, p. 9, pl. 6, igs 10–14, pl. 7, igs 7–9. 2008 Daphnogene cinnamomifolia (Brongniart in Cuvier 1822) Unger 1850; Winterscheid & Gregor, p. 4. 2002 M a t e r i a l. SMNS P 1952/ 8, 13, 15, 24 B, 38, 49, 51, 60, 64, 68, 71, 76, 77, 116, 121, 136, 140, 164, cf. 165, 174, 182, 185, 227, 228, 260, 261, 267, 273, 279, 296, 303, 314, 319, 320, 328, 331, 357, 358, 364, 415?, 418, 423, 425– 427, 430, 432, 449, 450, 451, 453, 455, 466, 471, 472; SMNS P 1953/1–14, 16–19, 21, 22, 49, 69, 76. D e s c r i p t i o n. Petiolate, entire-margined leaves, texture coriaceous, petiole up to 20 mm long, laminar shape oblong to elliptic or somewhat ovate, l × w = 19–110 × 7–47 mm, l/w ratio = 2.2–5.6, base straight to somewhat rounded, base angle acute, apex straight to acuminate, apex angle acute; midvein straight, Unangemeldet Heruntergeladen am | 10.04.17 15:37 339 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 basal secondaries suboppositely suprabasal acrodromous, usually prominent, running onto upper third of the lamina, further secondaries distinctly weaker, arising at angles of about 40–50° in broader leaves and narrower angles in smaller ones, such secondaries being restricted to the upper part of the lamina, brochidodromous, abmedially looping veinlets arising from suprabasal secondaries, tertiaries (mainly hardly visible) percurrent. M i c r o m o r p h o l o g y. Adaxial cuticle thick, anticlines of non-modiied cells distinct, from almost straight or curved to moderately undulate, cells 15–38 (20–31) µm across, largely glabrous to trichome bases rather densely spaced; abaxial cuticle delicate, in larger leaves more delicate than in smaller ones, anticlines of non-modiied cells delicate, rounded to undulate with smaller amplitude and wave length than adaxially, cells 12–35 (16–20) µm across, stomatal complexes brachyparacytic, oval to rhomboidal, often asymmetric, weakly staining, subsidiaries not offset from guard cells, aperture very short, slit-like, stoma length 10–20 (14–17) µm; trichome bases scattered to dense, considerably variable in shape and size, either situated in the plane of the cuticle or raised above the cuticle surface, trichome pore slit-like or a roundish pore 6–13 µm in diameter, sometimes with a thickened poral rim, encircling cells ± distinct, intensively staining and ± radially elongated; globular mesophyllous secretory bodies sometimes preserved. R e m a r k s. Variability of trichome density and of the undulation of the anticlines both adaxially and abaxially, largely going along with variability of leaf size, have been explained repeatedly as an expression of the leaf position on the living plant, that is, by whether they are sun or shade leaves, e.g., Kvaček & Walther (1978). Sometimes presumable shade leaves are described as forma lanceolata and sun leaves as forma cinnamomifolia, e.g., Walther (1999). Due to morphological overlaps, this distinction is not applied here. Trichome base variability (bases raised above surface or not, shape of encircling cells, shape and size of trichome pore, presence of distinct poral rim) has not yet been examined in terms of its taxonomic value. Specimen SMNS P 1953/76 is distinct due to the thick poral rim of the trichome pore (pl. 9, ig. 6), while the other cuticular features match Daphnogene very well. This specimen resembles Laurophyllum sp. 2 (Winterscheid & Kvaček 2014, pl. 9 ig. 12). Since the species diversity of Daphnogene-like foliage is still unresolved, the characteristics of the indumentum may help differentiate fossil species. Daphnogene cinnamomifolia is among the most common species in Rauenberg. Unlike at Flörsheim, no fragmentary twig has been recovered. The variability of laminar size is very similar in both localities (Flörsheim l × w = 50–120 × 10–53 mm, Rauenberg 19–110 × 7–47 mm). Laurus L. Laurus abchasica (Kolakovskii & Shakryl) Ferguson Pl. 6, igs 11–13; Pl. 10, igs 1–3 1958 Laurophyllum abchasicum, Kolakovskii & Shakryl in Kolakovskii, p. 346, pl. 7, igs 1–3, pl. 10, igs 4–8. 1974 Laurus abchasica (Kolakovskii & Shakryl) comb. nov.; Ferguson, p. 64, igs 7A–H, 8A–D, 9A–D, 10. 2004a Laurus abchasica (Kolakovskii & Shakryl) Ferguson; Kvaček, p. 6, pl. 4, igs 1, 2, pl. 5, igs 1–3. M a t e r i a l. SMNS P 1952/268, 377, 382; SMNS P 1953/65, 75. D e s c r i p t i o n. One almost complete leaf and fragments of narrow elliptic, entire-margined leaves, l × w up to 96 (about 120) × 22 mm, base ? acute, apex angle acute, apex shape almost straight to somewhat acuminate, texture moderately irm. M i c r o m o r p h o l o g y. Cuticles of both sides moderately thick, largely glabrous, anticlines of non-modiied epidermal cells strongly zig-zag undulate, undulation moderately deep, beadlike thickenings at tips of lobes, cells of similar size on both sides 15–44 (average 25–32) µm across, SMNS P 1953/65 with hypodermis adaxially; abaxial cuticle: stomatal complexes paracytic, shape roundish to rhomboidal, guard cells and subsidiary cells hardly staining and hardly delimited from each other, stomatal complexes l × w = 16–26 (average 18–21) µm × 13–28 (average 18–22) µm, l/w ratio = 0.6–1.5 (average 0.9–1.1), stomatal aperture spindleshaped, ledges slightly thickened, aperture length 5–16 (average 9–12) µm. R e m a r k s. A single leaf (SMNS P 1953/75) is Unangemeldet Heruntergeladen am | 10.04.17 15:37 340 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 almost complete. The largest leaf whose apical half is preserved probably measured about 110 mm in length and 22 mm in width. The cuticular structure of these remains clearly points towards Laurus abchasica. Laurophyllum ischkandelii Kunzmann et Walther (2002) is described from the late Eocene of the Weißelster Basin; it differs by its very narrow and strongly omega-like undulation of the anticlines both adaxially and abaxially. Remarkable is the presence of a hypodermis on specimen SMNS P 1953/65 from Rauenberg. Ferguson (1971) describes for taxon XXIII from Kreuzau, which corresponds to Laurus abchasica, “primary cells often subdivided by secondary cell walls” on the adaxial cuticle, but does not mention whether this refers to a hypodermis. L. abchasica records are more common during the late Oligocene and the subsequent Miocene than in the early Oligocene. Apart from two leaves recorded from Flörsheim (Kvaček 2004a), Rauenberg is the second early Oligocene record. Laurophyllum Goeppert Laurophyllum kinkelinii (Engelhardt) Kvaček stomata overlapped by subsidiaries, stomatal aperture narrow spindle-shaped to narrow elliptic to slit-like, ledges not very pronounced, stoma length 16–25 (18, 19) µm, trichome bases common, indistinct; mesophyllous secretory bodies rare. R e m a r k s. While specimens SMNS P 1952/275+276 and SMNS P 1953/64 are fragments of large, probably narrow elliptic leaves, SMNS P 1952/184 is a long, narrow elongate fragment. These remains differ from the below-described Laurophyllum rauenbergense by their bigger leaf size, less coriacous lamina, distinctly bigger cells of the adaxial cuticle and dense spacing of the trichome bases abaxially. Unlike L. kinkelinii from Flörsheim, these specimens lack dense mesophyllous secretory bodies. It remains open whether this is an artefact caused by preservation or cuticle preparation. The close similarities to L. saxonicum Litke and L. medimontanum Kvaček have been pointed out by Kvaček (2004a), as was a possible relationship of L. kinkelinii to Persea. Laurophyllum cf. kinkelinii (Engelhardt) Kvaček 2004 var. glabrum Kvaček Pl. 2, igs 7, 8; Pl. 11, igs 5–9 Pl. 2, igs 9–11; Pl. 12, igs 1–4 2004a Laurophyllum kinkelinii (Engelhardt) Kvaček comb. n.; Kvaček, p. 7, pl. 4, igs 10–12, pl. 5, igs 10–12. 2004a Laurophyllum kinkelinii (Engelhardt) Kvaček comb. n. var. glabrum Kvaček var. n.; Kvaček, p. 7, pl. 4, igs 13–16, pl. 7, igs 1–3. M a t e r i a l. SMNS P 1952/184, 275+276 (part and counterpart); SMNS P 1953/64. M a t e r i a l. SMNS P 1952/378, 396; SMNS P 1953/87. D e s c r i p t i o n. Entire-margined leaves moderately coriaceous, petiole up to 15 mm long at least, laminar shape (narrow) elliptic, oblong, l × w = up to at least 120 × 18–25 mm, l/w ratio about 4.8–6.7, leaf base narrow acute, apex not preserved; midvein stout, mainly straight, secondaries vaguely visible, rather widely spaced, curved. D e s c r i p t i o n. Macromorphology very similar to L. kinkelinii; l × w = 53 (? 65)–62 (? 90) × 15–19 mm, l/w ratio about 3.2–6.7. M i c r o m o r p h o l o g y. Adaxial cuticle thick, well preserved, anticlines distinct, smooth, straight, curved to shallow undulate with wide wave length, non-modiied cells 19–41 (31, 31) µm across, largely glabrous, epicuticular striae mainly above veins and near margin, radial striae around trichome bases; abaxial cuticle delicate, non-modiied cells domed, anticlines therefore often dificult to trace, 12–24 (18, 21) µm across, stomatal complexes paracytic, Micromorphology also very similar to L. kinkelinii except for: adaxially, anticlines slightly undulate, non-modiied cells 22–45 (33–37) µm across, striation upon veins and radial striae near rare trichome bases; abaxially scattered large trichome bases with distinct radial striation; mesophyllous secretory bodies present. R e m a r k s. SMNS P 1952/396 and SMNS P 1953/87 are medium-sized, elliptic specimens (l/w ratio about 3.2 and 4), while specimen SMNS P 1952/378 is somewhat larger and more elongated (l/w ratio about 6.7). These leaves show larger cells with somewhat undulate anticlines on the adaxial cuticle as compared to L. kinkelinii. They differ from the Flörsheim Unangemeldet Heruntergeladen am | 10.04.17 15:37 341 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 material by the striae adaxially and abaxially and the lower frequency of mesophyllous secretory cells. Laurophyllum pseudoprinceps Weyland & Kilpper s.l. Pl. 2, igs 12–16; Pl. 12, igs 5–7 Laurophyllum pseudoprinceps n. sp.; Weyland & Kilpper, p. 23, igs 14–19, textig. 6. 1971b Laurophyllum pseudoprinceps Weyland & Kilpper; Kvaček, p. 50, pl. 1, igs 4–6, pl. 3, igs 3–5, pl. 4, igs 1–4, text-ig. 1. 2004a Laurophyllum pseudoprinceps Weyland & Kilpper s.l.; Kvaček, p. 8, pl. 4 igs 3–5, pl. 5, igs 7–9. 1963 M a t e r i a l. SMNS P 1952/22+27, 30+31, ? 217, 265, 270, 282, 302, 312, 344, 345, 362, 365, 389, 392, 403, 409, 412, 446; SMNS P 1953/15A, 48, 84. D e s c r i p t i o n. Fragments of coriaceous leaves, rarely almost complete, laminar shape elliptic to narrow elliptic, l × w = 40–85 × 10–28 mm, l/w ratio = 2.3–4.3, base angle (narrow) acute, base mainly cuneate, sometimes slightly convex or wavy, petiole complete in specimen SMNS P 1952/312: 10 mm long, straight, widened at base; apex angle (narrow) acute, apex mainly straight to somewhat acuminate, margin entire, sometimes line-like thickened, midvein straight, secondaries only rarely visible, basal ones sometimes more prominent. M i c r o m o r p h o l o g y. Adaxial and abaxial cuticles thick, largely glabrous, anticlines variable from almost straight to undulate, often pitted, especially adaxially; non-modiied epidermal cells 15–32 (average 18–24) µm across adaxially; abaxial cuticle: non-modiied epidermal cells 16–36 (average 23–30) µm across, stomatal complexes paracytic, often asymmetric, butterly-like in shape due to the presence of big lateral subsidiary cells bordering the sunken guard cells, stomatal ledges thick, stomata length 17–27 (average 20–23) µm, specimen SMNS P 1953/15 stoma length 20–33 (average 24) µm; mesophyllous oil cells present. R e m a r k s. The stoma length of the type material of Laurophyllum pseudoprinceps is indicated as “meist zwischen 17 und 21 µ” (Weyland & Kilpper 1963). Kvaček (1971b) characterises the stoma length of L. pseudoprinceps as 17–25 µm. Ocotea hradekensis resembles L. pseudoprinceps strongly except for the larger stomata 25–30 (–35) µm (Kvaček & Bůžek 1966, Bůžek et al. 1996). This species has also been reported from Markvartice although without measurements (Bůžek et al. 1976), from Hradek nad Nísou (Holý et al. 2012), from the Cypris Shale (Bůžek et al. 1996) and from the Oder 2a lignite mine (Knobloch & Kvaček 1976) with the following variability in stoma length: 25–30 (36) µm (Hradek), (20–) 23–27 (–30) µm (Cypris Shale), and (22–) 27–30 (34) (Oder 2a). For L. pseudoprinceps from Flörsheim, Kvaček (2004a) describes the stomata as 22–25 (–30) µm long. This brief enumeration points to the dificulties of separating the two species based on stoma length. All specimens from Rauenberg included in L. pseudoprinceps bear stomata with an average length from 20–23 µm, larger than that described for the type material of L. pseudoprinceps by Weyland & Kilpper (1963). However, they well match the stoma size of L. pseudoprinceps given by Kvaček (1971b). Only in specimen SMNS P 1953/15 are the stomata somewhat longer: 20–33 µm (average length 24 µm). Thus it remains disputable whether this single specimen should be better assigned to Ocotea hradekensis. Laurophyllum rauenbergense Kovar-Eder sp. nov. Pl. 2, igs 17–25; Pl. 10, igs 4–9 2004a Laurophyllum cf. villense (Weyland & Kilpper) Kvaček; Kvaček, p. 8, pl. 6, igs 1–7, pl. 7, igs 4–6. Holotype 1953/82. designated h e r e. SMNS P P a r a t y p e s. SMNS P 1952/93, 391, 445; SMNS P 1953/47, 73, 85. S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. Locus t y p i c u s. Rauenberg, Württemberg, Germany. Baden- D e r i v a t i o n o m i n i s. Referring to the town of Rauenberg, where the fossil outcrop is located. A d d i t i o n a l m a t e r i a l. SMNS P 1952/278, 350, 379, 395, 397. Cf.: SMNS P 1952/231, P 1953/59. Unangemeldet Heruntergeladen am | 10.04.17 15:37 342 Based on gross morphology SMNS P 1953/91,? 94. J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 o n l y. D i f f e r e n t i a l d i a g n o s i s. Differing from L. villense (Weyland & Kilpper) Kvaček by its short stomata and absence of striae, and from L. kinkelinii by its smaller, non-modiied epidermal cells of the adaxial cuticle. D e s c r i p t i o n. Rather small, narrow, entiremargined, coriaceous leaves; petiole up to 16 mm long, laminar shape oblong to somewhat ovate or minimally elliptic, l × w = 35–72– ? 120 × 7–10 mm, exceptionally up to 13 mm, l/w ratio = 4.4–9.2, base cuneate to decurrent, base angle narrow acute, base straight to somewhat convex, midvein stout, mainly straight, secondaries not visible. M i c r o m o r p h o l o g y. Adaxial cuticle: thickness variable, moderately thick or thick, anticlines slender to thick, sometimes knobby, mostly straight to curved, non-modiied cells 9–30 (13–24) µm across, in specimens SMNS P 1952/391 and SMNS P 1953/82 domed, mainly glabrous; abaxial cuticle: delicate, nonmodiied cells domed or almost undomed, often dificult to trace, 11–33 (19–27) µm across, stomatal complexes paracytic, stoma shape variable due to subsidiaries which partly cover guard cells, aperture narrow, slit-like, stomatal ledges unpronounced, stoma length 12–21 (14–18) µm, trichome bases indistinct, rare to almost glabrous; mesophyllous secretory bodies present. R e m a r k s. This Lauraceae is characterised by its small, linear to narrow ovate and oblong leaves. The length/width ratio of the lamina is usually >5. In Flörsheim this laurel is even more common than Daphnogene foliage. Kvaček (2004a) hesitated about the unambiguous assignment to Laurophyllum villense, probably due to the cuticular striation (Weyland & Kilpper 1963) and the younger age of the type material of L. villense. More relevant is the difference in stoma length, which is shorter and very similar in Flörsheim (16– 20 µm, Kvaček 2004a) and Rauenberg (12–21 (average 14–18) µm) as compared to L. villense (up to 28 µm, Weyland & Kilpper 1963). The assignment of the specimens SMNS P 1952/231 and SMNS P 1953/59, which match well the gross morphology of L. rauenbergense, is somewhat ambiguous because the stomata are less masked by the subsidiary cells. Furthermore, specimen SMNS P 1952/231 bears rare trichome bases both adaxially and abaxially, with faint radial striation around the trichome base, which could point towards L. kinkelinii var. glabrum (see above). In the latter species, however, the cells of the adaxial epidermis are distinctly bigger than in L. rauenbergense. Moreover, leaves larger than typical L. rauenbergense forms are sometimes dificult to assign unambiguously to either L. rauenbergense or L. kinkelinii (see also Kvaček 2004a). Laurophyllum sp. A Pl. 6, ig. 8; Pl. 12, igs 8, 9 M a t e r i a l. SMNS P 1952/336. D e s c r i p t i o n. Almost complete, entire-margined leaf, laminar shape elliptic, l × w = 67 (? 72) × 29 mm, l/w ratio about 2.5, base angle acute, base almost straight to weakly ? obtuse, apex angle acute, apex acuminate, secondaries prominently (supra)basal acrodromous. M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle thick, non-modiied epidermal cells on both sides very similar, anticlines straight, thick, cell size adaxially 15–22 (average 18) µm, abaxially 14–26 (average 20) µm; abaxial cuticle: stomatal complexes paracytic, subsidiary cells resembling the non-modiied epidermal cells in shape and size, stoma length 12–18 (average 15) µm, (stoma width unclear), aperture indicated by short, thickened ledges, aperture length 5–10 (average 7) µm; mesophyllous oil cells present. R e m a r k s. Based on gross morphology this leaf coincides with Daphnogene cinnamomifolia. It differs distinctly by having a thick abaxial cuticle and prominent anticlines. The stomatal complexes are neither rhomboidal nor asymmetric, and they stain as well as the other cells. The assignment among the Lauraceae is supported by mesophyllous oil cells. Laurophyllum sp. B Pl. 6, ig. 9; Pl. 13, igs 1–3 M a t e r i a l. SMNS P 1952/47. D e s c r i p t i o n. Basal part of a coriaceous leaf, base angle acute, base probably decurrent; petiole 14 mm long, fragmentary, lamina Unangemeldet Heruntergeladen am | 10.04.17 15:37 343 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 l × w = 70 (? 140) × 15 (?) mm, midvein distinctly bent, base probably entire-margined. M i c r o m o r p h o l o g y: Adaxial and abaxial cuticle well preserved, both distinctly granular, adaxial one thicker than abaxial one, anticlines straight to somewhat curved, thicker adaxially than abaxially, non-modiied cells 18–31 (abaxially 26, adaxially 24) µm across, glabrous; large mesophyllous oil glands present, e.g., 34 µm in diameter; abaxial cuticle: stomatal complexes brachyparacytic, subsidiary cells sickle-shaped, not or hardly offset from guard cells, guard cells sunken, aperture slit-like, ledges slender lines, stoma length 11–23 (18) µm, aperture length 6–12 (9) µm. R e m a r k s. This leaf may belong to Daphnogene because it is possible that the suprabasal acrodromous secondaries are masked due to fragmentary preservation. The stomatal complexes resemble those of Daphnogene but are relatively well cutinised and larger. The relatively robust abaxial cuticle and the granular structure of both surfaces are further details pointing towards a different systematic position within the Lauraceae. Laurophyllum sp. C Pl. 6, ig. 10; Pl. 13, ig. 4 M a t e r i a l. SMNS P 1953/81. D e s c r i p t i o n. Narrow oblong, entire-margined leaf, texture irm, l × w = 65 × 8 mm, l/w ratio = 5.9, base and apex attenuate, petiole 5 mm (incomplete), curved; midvein straight, secondaries weakly brochidodromous, widely spaced, ascending under steep angles (about 30°) and somewhat bent. M i c r o m o r p h o l o g y. Adaxial cuticle well cutinised, anticlines slender, smooth, widely undulate, non-modiied cells 16–48 (average 29) µm across; abaxial cuticle poorly preserved, only stomata discernable, stomata sunken, 13–16 (average 15) µm long, only the area close to aperture visible, slender ledges marking the narrow aperture; both sides glabrous. R e m a r k s. There is no doubt about the family assignment of this leaf. Gross morphologically it resembles Laurophyllum rauenbergense, but the adaxial cuticle is more reminiscent of L. kinkelinii. The stomata are small even for L. rauenbergense. Arecales Arecaceae Schultz Palmacites Brongniart Palmacites lamanonis Brongniart Pl. 3, igs 1, 2; Pl. 13, ig. 5 1822 Palmacites lamanonis, Brongniart, p. 38, pl. 3, ig. 1. 2004a Palmacites lamanonis Brongniart; Kvaček, p. 23, pl. 20, igs 1, 2. 2008 Trachycarpus (Flabellaria) raphifolia (Sternberg) Takhtajan; Winterscheid & Gregor, pl. 2, ig. 2. M a t e r i a l. SMNS P 1952/66, 74, 75, 95, 96, 111, 112, 113, ? 122, 129, 175, 280; SMNS P 1953/113+114, 124. D e s c r i p t i o n. Fragments of fan-shaped leaves, lamina plicate, petiole unarmed, hardly extending into the lamina; specimen SMNS P 1953/124 most complete, petiole l × w = 112 × 33 mm (incomplete), fragment of lamina 225 mm long, 20 mm wide. M i c r o m o r p h o l o g y. Cuticle moderately coriaceous, ? hypostomatic, adaxial cuticle: anticlines irregularly thick, areas with irregularely shaped cells, 12–25 (average 16) µm across, alternating with narrow elongated cells with striae; abaxial cuticle in stoma-bearing condition: anticlines regularly slender, straight to somewhat curved, cells ± elongated to shorter than wide, cross walls mainly oblique, l × w = 10–15 (average 13) × 5–11 (average 7) µm, stomatal complexes loosely spaced, brachyparatetracytic, guard cells often indistinct and masked by fungi, ledges thickened, stomata l × w = 14–20 × 10–18 µm (n = 3). R e m a r k s. According to the pragmatic classiication provided by Read and Hickey (1972), these leaves match the fossil genus Palmacites, unlike those described below. The cuticle does not allow further systematic assignment (compare Mai & Walther 1978). Fragments of fan palms lacking a costa are more common in Rauenberg than in Flörsheim (Kvaček 2004a). Sabalites Saporta Sabalites major (Unger) Saporta Pl. 3, igs 3, 4 1847 Flabellaria major, Unger, p. 42, pl. 14, ig. 2. 1865 Sabalites major (F. Unger) G. Saporta, p. 83, pl. 2. Unangemeldet Heruntergeladen am | 10.04.17 15:37 344 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 M a t e r i a l. SMNS P 1952/23, 132, 169, 173; SMNS P 1953/115. D e s c r i p t i o n. Fragments of costapalmate leaves, lamina plicate, petiole unarmed, hastula acuminate-attenuate; most complete specimen (SMNS P 1952/169): petiole incomplete, l × w = 105 mm × 35 mm, fragment of fan-shaped lamina 225 mm long, 145 mm wide, hastula extending about 100 mm into the lamina. R e m a r k s. These palm leaves differ from those described above by having the hastula extending considerably into the lamina. In modern palms, hastulas may be very differently expressed adaxially and abaxially in one and the same species (Dransield et al. 2008). Therefore it cannot be ruled out that these leaves represent the abaxial side of the above-described taxon Palmacites lamanonis. Phoenicites Brongniart Phoenicites sp. Pl. 3, igs 7–9; Pl. 5, ig. 1; Pl. 13, ig. 9 2004a ? Phoenicites sp.; Kvaček, p. 23, pl. 20, igs 3, 4. 2008 Phoenicites sp.; Winterscheid & Gregor, pl. 2, ig. 1. M a t e r i a l. SMNS P 1952/12, 141, 473; SMNS P 1953/116. SMNK 7793. NMA 2006-112/1497, 2006-180/1497. D e s c r i p t i o n. Fragments of pinnately dissected foliage; pinnae suboppositely arranged, rhachis up to 11 mm wide; most complete fragment of a rhachis 178 mm long (NMA 2006112/1497), pinnae plicate, narrowing into cuneate base, divided into two unequally wide segments, longest pinna fragment 140 mm, venation very ine, densely parallel. M i c r o m o r p h o l o g y. Cuticle very delicate, cell outlines often indistinct, in non-stomatal regions anticlines straight or somewhat curved, otherwise shallow undulate, cell outlines rectangular to oblique, arranged in rows, 12–16 µm long and 5–7 µm wide (n=6); stomatal complexes in bands, rather widely spaced, oriented parallel to the pinna axis, paratetracytic, number of lateral subsidiary cells obscure, guard cells weakly cutinised, stomata small, 14–22 (average 16) µm long, front cavity wide elliptic, 8–10 µm long (n=5), cuticular ledges distinct, their ends not meeting each other, aperture spindle-shaped. R e m a r k s. The most complete specimen is that igured by Winterscheid and Gregor (2008: pl. 2 ig. 1), which is on exhibit in NMA. Other more complete specimens (SMNS P 1953/116, NMA 2006-112/1497) have been embedded by transfer technique in artiicial resin. Although this hampers cuticle studies, the method prevented the specimens from falling apart. The other specimens are far less complete. SMNS P 1952/12 is an axis with the very basal parts of the pinnae preserved only; it is the only one with cuticle preservation. In SMNS P 1952/141, the rhachis is more slender and the subdivision of the pinnae is well visible. The material at hand does not allow determination of whether the segmentation of the pinnae is of diagnostic value. A single similar specimen has also been described from Flörsheim (Kvaček 2004a). Though the segmentation of the pinnae is not mentioned there, the close-up view in pl. 20 ig. 4 resembles our specimens. Phoenicites borealis Friedrich specimens from Geiseltal (Rüfle 1976) and the Weisselster Basin (Mai & Walther 1985) differ by having twice-bigger stomata and by their thicker cuticular ledges forming the front cavity. ? Calamoideae gen. et sp. indet. Pl. 3, ig. 6; Pl. 13, igs 6–7a M a t e r i a l. NMA 2016-1 /1497. D e s c r i p t i o n. Fragment of a very strongly armed leaf or petiole, length 108 mm (incomplete), width 30 mm (incomplete); hook-shaped, very massive spines regularly spaced, attached to the ? rhachis or ? petiole, spines 3–4 mm long and up to 3 mm wide at their base; venation of adjacent tissue parallel with perpendicular anastomoses. M i c r o m o r p h o l o g y. Non-modiied cells ± distinctly arranged in rows paralleling the length of the ? leaf, but cells not elongated, 7–15 (average 12) µm long, rectangular to oblique, anticlines thick, appearing in double lines; stomatal complexes loosely spaced, paratetracytic to cyclocytic, oriented mainly paralleling leaf length, guard cells sunken, stomatal aperture narrow, spindle-shaped, with distinct cuticular ledges, stoma length 12–19 (average 15) µm, aperture length 8–11 (average 9) µm. R e m a r k s. This remarkable remain is embedded in artiicial resin. It deinitely derives Unangemeldet Heruntergeladen am | 10.04.17 15:37 345 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 from a palm. This is suggested by the parallel venation in combination with the spines and conirmed by the cuticles, i.e., stomatal complexes. Nevertheless, it is somewhat dificult to interpret which part of the leaf is at hand. The regularly spaced spines are attached to one side of a thick, possibly even woody rhachis or petiole, whereby on the other side a fragment of the lamina appears attached. Armature is quite common among palms, and not restricted to Calamoideae (Dransield et al. 2008). Arecaceae gen. et sp. indet. Pl. 3, ig. 5; Pl. 13, igs 7b–8 2008 Problematicum: cf. Arecaceae gen. indet.; Winterscheid & Gregor, p. 6, pl. 3 ig. 1. M a t e r i a l. NMA 2016–2/1497. D e s c r i p t i o n. Probably a leaf, petiole stout, length 37 cm, incomplete, width 8 cm, ? lamina consisting of ine, lexible iliform segments, segments up to about 23 cm long, of uncertain arrangement. M i c r o m o r p h o l o g y. Cells arranged in rows, not clearly traceable, stomata scattered, arranged parallel to cell rows, 36–41 µm long, stomatal complexes probably paratetracyctic. R e m a r k s. This specimen is characterised by the peculiar structure of its lamina. The cuticle points towards afinity to Arecaceae. The stomata are distinctly larger than in Phoenicites sp. and even larger than in P. borealis (Rüfle 1976, Mai & Walther 1985). This specimen was published by Winterscheid and Gregor (2008) erronously under coll. no. 2006-181/1497. Proteales Platanaceae T. Lestib. Platanus L. Platanus neptuni (Ettingshausen) Bůžek, Holý et Kvaček Pl. 4, igs 1–10; Pl. 14, igs 1–6 1967 Platanus neptuni (Ettingshausen 1866) comb. nov.; Bůžek et al., p. 203, pls 1–4. 2004a Platanus neptuni (Ettingshausen) Bůžek, Holý et Kvaček; Kvaček, p. 10, pl. 8, igs 1–13. M a t e r i a l. SMNS P 1952/ 3, 4, 5, 9, 14, 16, 19, 24 A, 25, 26, 32+97 (part and counterpart), 37, 42, 59, 81+84, 82, 86, cf. 128, 130, 133, 137, 151, 153, 160, 178+179 (part and counterpart), 180, 181, 195, 201, 202, 212, 216, 218, 219, 220, 221, 230, 236, 239, 241, 253, 257, 263, 266, 272, 277, 297, 304, 308, 309, 341, 346, 352, 355, 363, 387, 398, 401, 424, 429, 433, 434, 435, 436, 438, 439, 440, 443+448 (part and counterpart), 462, 464, 465, 467, 469, 474; SMNS P 1953/20, 23, 24A–32, 34, 36–42, 44–46, 67, 78, 83, 96, 98, 120. SMNK 7480, 7598 left, 7664, 7669, 7688, 7727, 7812, 7891. NMA 2006-117/1497, 2006120/1497, 2006-128, 129/1497, 2006-148/1497, 2006-151/1497, 2006-153/1497, 2006-155/2006, 2006-158/1497, 2006-161/1497. D e s c r i p t i o n. Simple petiolate leaves, texture coriaceous, lamina oblong, narrow elliptic, mostly symmetric but sometimes also somewhat asymmetric, variable in size, l × w = 29 (38)–160 (180) × 15–73 mm, l/w ratio = 2–4.4; petiole up to 15 mm long, basally widened; base angle acute to obtuse, base straight or somewhat concave to convex, sometimes cuneate to decurrent, apex angle acute, apex straight to acuminate, leaf margin mostly entire near base of the lamina, then simple serrate, sometimes generally almost entire; teeth mainly small with long basal and very short apical sides, apex mainly rounded, sometimes even hook-shaped, densely or widely spaced but at rather regular distances within single leaves, some leaves rather coarsely serrate; midvein mostly straight, secondaries brochidodromous, rather widely spaced, intersecondaries present; further venation details available only from specimen NMA 2006-128/1497: secondaries arching with apically adjacent secondary; marginally, further smaller-scale arches are formed by veinlets originating from secondaries, free veinlets end up in tooth apices of the margin; tertiaries and fourth-order venation random reticulate. M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle thick, mostly well preserved; anticlines of non-modiied epidermal cells slender, mostly distinct, undulated, undulation U-shaped, wide, giving the appearance of puzzle-like indenting, or small and narrow, non-modiied epidermal cells 30–70 (average 37–56) µm adaxially and 22–67 (average 34–47) µm abaxially across, trichome bases mainly two- to four-celled, single-celled mainly upon veins, periphery strongly thickened, adaxially rare to almost absent, abaxially scattered or more abundant, Unangemeldet Heruntergeladen am | 10.04.17 15:37 346 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 inner diameter 12–25 µm, attached peltate trichomes rare, 46–73 µm in diameter, only exceptionally cellular partitioning visible, cuticule surface striate to wrinkled, parallel upon veins and near leaf margin, sometimes present also in non-venous areas, striation stronger and running radially around trichome bases; abaxial cuticle: stomatal complexes anomocytic, irregularly and widely spaced, stomata elliptic, l × w = 27–45 (average 32–40) × 22–38 (average 27–34) µm, l/w ratio = 0.9–1.4 (average 1.1–1.2), epidermal wall of guard cells thickened, ledges thickened, forming an elliptic to broad spindle-shaped outer front cavity, 14–36 (average 18–31) µm long, aperture a ± faint slit. R e m a r k s. This is by far the most common fossil species at Rauenberg, accounting for about one sixth of all plant remains (>100). Most leaves are symmetric and no compound specimen was recovered. Therefore, we are dealing with leaves of P. neptuni rather than with P. fraxinifolia-type lealets. While most specimens can be determined based on gross morphology, some are more ambiguous; some such specimens in the museum´s collection were found determined as, for example, Carya (P 1952/32+97, part and counterpart, pl. 4, ig. 1). Based on the cuticles, the assignment to Platanus neptuni is unambiguous. Although foliage is abundant, no fruiting heads of Platanus were detected. Buxales ? Buxaceae Dumort. ? Buxus L. ? Buxus egeriana Kvaček, Bůžek et Holý Pl. 6, ig. 19 M a t e r i a l. NMA 2006-168/1497. D e s c r i p t i o n. Coriaceous, entire-margined leaf, petiole fragmentary, 1 mm long, laminar shape narrow elliptic to oblong, l × w = 27 (28) × 8 mm, l/w ratio = 3.5, base angle acute, base straight, apex ? emarginate, midvein stout, straight, secondaries very densely spaced, probably brochidodromous, single secondaries forking, further venation not discernable, margin strongly thickened, probably by a imbrial vein. M i c r o m o r p h o l o g y. Cuticle poorly preserved, anticlines of non-modiied cells straight, forming vaguely traceable polygonal cell outlines, stomatal complexes ? anomocytic, roundish with broad spindle-shaped aperture (one well visible, others masked by organic material). R e m a r k s. This leaf was transfered to artiicial resin. The organic material is preserved only fragmentarily but the secondary venation is discernable on the resin surface. The assignment remains somewhat ambiguous because the ramiication of the venation characteristic of Buxus is not preserved and due to poor cuticle preservation. Certiication of this record would extend the stratigraphic range of Buxus egeriana, which was known hitherto from the earliest Miocene (Linz, Austria, Kovar 1982, Cypris Formation, Kvaček et al. 1982). Fagales Fagaceae Dumort. Eotrigonobalanus Walther & Kvaček Eotrigonobalanus furcinervis (Rossmässler) Walther & Kvaček forma haselbachensis (Kvaček & Walther) Kvaček & Walther in Walther Pl. 6, ig. 14; Pl. 14, ig. 7 1989a Dryophyllum furcinerve (Rossmässler) Schmalhausen forma haselbachensis forma nova; Kvaček & Walther, p. 220, igs 2 d, 3 a, c. 1989b Eotrigonobalanus furcinervis (Rossmässler) Walther & Kvaček forma haselbachensis (Kvaček & Walther) comb. nov.; Kvaček & Walther, p. 584, igs 7–10, pls 34–36, pl. 39, igs 2–4, pls 44–46. 2004a Eotrigonobalanus furcinervis (Rossmässler) Walther & Kvaček ssp. haselbachensis (Kvaček & Walther) Kvaček & Walther in Walther; Kvaček, p. 13, pl. 10, igs 10, 11, 17. Non 2008 Eotrigonobalanus furcinervis (Rossmässler) Walther & Kvaček 1989; Winterscheid & Gregor, pl. 1 ig. 4. M a t e r i a l. SMNS P 1953/60. D e s c r i p t i o n. Single petiolate leaf, lamina oblong, slightly obovate, l × w = 85 (95–100) ×22 mm, l/w ratio about 4.4, base angle acute, base cuneate running into the thick petiole, petiole length 5 mm, incomplete, margin entire, midvein straight, stout, secondaries Unangemeldet Heruntergeladen am | 10.04.17 15:37 347 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Betulaceae Gray vel Ulmaceae Mirbel slender, arising at about 45° and rather regular distances, gently curved, presumably brochidodromous. M i c r o m o r p h o l o g y. Adaxial cuticle not preserved; abaxial cuticle delicate, only above veins anticlines rather well discernable, straight, stomatal complexes in groups, densely spaced, ? cyclocytic, shape roundish, stomata l × w = 19–25 (average 22) × 20–23 (average 22) µm, l/w ratio = 0.9–1.2 (average 1), cuticle above guard cells thin, stomatal ledges indistinct, front cavity short elliptic to roundish, trichome bases scattered, 6–11 (9 average) µm in diameter. R e m a r k s. Remarkably, this is the only leaf representing the Fagaceae in Rauenberg. Although the cuticle is not very well preserved, its systematic afinity is clear due to the combination of gross morphological and cuticular features. This mainly entire-margined subspecies largely devoid of trichome bases of stellate trichomes is rather common in the late Eocene and early Oligocene. Due to the cuneate leaf base, cf. Trigonobalanopsis sp. (Mallison 2002, pl. 10 ig. 1) from the Alzey Formation in the Mainz Basin may represent a leaf of Eotrigonobalanus rather than of Trigonobalanopsis. Betulaceae Gray Betula L. Betula dryadum Brongniart Betulaceae vel Ulmaceae gen. et sp. indet. Pl. 6, ig. 20; Pl. 14, ig. 8 M a t e r i a l. SMNS P 1952/437. D e s c r i p t i o n. Petiolate, membranaceous leaf, petiole slender, 4 mm long, incomplete; lamina asymmetric, ovate, l × w = 56 (? 62–65) × 20 mm, l/w ratio about 3.1–3.3, base angle obtuse, base rounded, apex angle acute, apex shape incomplete, probably straight; margin sharply and densely serrate, probably double serrate; midvein slender, straight, secondaries delicate, somewhat bent, spacing of secondaries 3–5 mm. M i c r o m o r p h o l o g y. Cuticle delicate, anticlines slender, straight to somewhat bent, cell outlines polygonal, 17–22 µm across, ? stomata indicated as more strongly cutinised, broad oval spots, ? anomocytic. R e m a r k s. Gross morphologically, this leaf is reminiscent of Betulaceae and, due to the asymmetric shape, afinity to Ulmaceae cannot be excluded. It remains open whether the scattered thickened regions on the cuticle are interpreted correctly as stomata. From Flörsheim a single leaf ascribed to Ostrya atlantidis Unger (Kvaček 2004a) differs by having marginal serration which is distinctly double serrate, with irst-order teeth much longer than the second-order ones. Pl. 6, ig. 1 Myricaceae Rich. ex Kunth 1828 Betula dryadum Brongniart, p. 49, pl. 3, ig. 5. Comptonia L. Hérit. M a t e r i a l. SMNK 7711. D e s c r i p t i o n. Two-winged fruit, nutlet narrow obovate, 3.2 mm long, 1 mm wide at widest part, style remnant incomplete, wings skin-like, fruit width including wings about 4.5 mm. R e m a r k s. Fruits of this type are usually assigned to Betula dryadum. Often they are found associated with Betula bracts as in Armissan (Upper Oligocene, France; Brongniart 1828) or Seifhennersdorf (Lower Oligocene, Germany; Mai 1963). From Rauenberg, however, no bracts have been detected so far, although they are woody and therefore predestined for fossilsation. Comptonia difformis (Sternberg) Berry Pl. 6, igs 6, 7 1821 Asplenium difforme, Sternberg, p. 29, 33, pl. 24 ig. 1. 1825 Aspleniopteris difformis Sternberg, p. 21, pl. 24, ig. 1. 1906 Comptonia difforme (Sternb.); Berry, 495. 2004a Comptonia schrankii (Sternberg) Berry; Kvaček, p. 12, pl. 9, ig. 13. M a t e r i a l. SMNS P 1952/166, SMNS P 1953/118. NMA 2006-146/1497. D e s c r i p t i o n. Pinnate leaves, SMNS P 1952/166 35 mm long (about 2/3 to 3/4 of Unangemeldet Heruntergeladen am | 10.04.17 15:37 348 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 complete length preserved), 3.5 mm wide, SMNS P 1953/118 57 mm long, ? half of complete length, 8 mm wide, pinnae tightly spaced, basal side convex, apical side straight to gently concave, attached with full width to midvein, pinnae widest in the middle part of the leaf, tapering towards base and apex; in specimen SMNS P 1953/118, pinnae and especially their margin slightly curled; venation of pinnae not visible. R e m a r k s. Specimen SMNS P 1952/166 better matches Comptonia schrankii (Sternberg) Berry but the other one (SMNS P 1953/118) more closely resembles C. difformis (Sternberg) Berry. Kvaček (2004a) supposes that both species relect ecotypes of a single species rather than discrete species; this is supported by the co-occurrence of the two types in Rauenberg. Myrica L. Myrica cf. lignitum (Unger) Saporta sensu Ettingshausen & Standfest Pl. 4, ig. 11; Pl. 14, ig. 9 Quercus lignitum Ung.; Unger, p. 113, pl. 31, igs 5–7. 2004a Myrica lignitum (Unger) Saporta; Kvaček, p. 11, pl. 9, igs 11, 12, 16, 17. roundish, weakly cutinised, aperture broad roundish, acute at poles, l × w=20–25 (average 23) × 20–23 (? 27) (average 21) µm, aperture length 9–12 (average 10) µm. R e m a r k s. The wide variability of Myrica lignitum leaves is well known not only from the type locality of Parschlug, Austria (Miocene; Kovar-Eder et al. 2004), but also from the marine deposits of the surroundings of Linz, Austria, earliest Miocene (Kovar 1982). The here- described specimen is the only one from Rauenberg that meets M. lignitum gross morphologically. Due to the scarcity of peltate trichomes as well as the thinly cutinised anticline separating the base cells of the trichome bases, the assignment remains a bit ambiguous. Recently, Winterscheid and Kvaček (2014) igured a specimen as M. lignitum from Orsberg near Bonn, discussing its gross morphological similarity to M. joannis (Ettingshausen) emend. Kovar-Eder. The acute teeth starting already near the leaf base support the assignment to M. joannis instead of M. lignitum. 1847 M a t e r i a l. SMNS P 1952/361. D e s c r i p t i o n. Moderately coriaceous leaf, shape somewhat obovate, l × w = 60 (? 65) ×16 mm, l/w ratio about 4.1, base cuneate, petiole not preserved, apex incomplete, ? acuminate, margin entire in the lower part of the lamina, then simple serrate, teeth distinctly developed, hook-shaped, basal side convex, apical side concave, sinus rounded, apex blunt to acute; midvein almost straight, secondaries only exceptionally and faintly traceable, curved upwards eucamptodromous or brochidodromous. M i c r o m o p h o l o g y. Cuticle delicate, adaxially and abaxially anticlines straight to curved, distinct, non-modiied cells 29–40 (average 35) µm across adaxially, 19–26 (average 23) µm abaxially; trichome bases extremely rare, roundish, bicellular, joint anticline of base cells weakly but margin strongly cutinised, inner diameter 10–12 µm, a single peltate trichome observed, 76 µm across; abaxial cuticle: stomatal complexes anomocytic, stomata Myrica longifolia Unger Pl. 4, igs 18–20; Pl. 15, igs 1, 2 1850 Myrica longifolia, Unger, p. 159, Taf. 27 Fig. 2, Taf. 28 Fig. 1. 2004a Myrica longifolia Unger; Kvaček, p. 11, pl. 9, igs 5–8, 14, 15. 2008 Myrica lignitum (Unger 1847) Saporta 1865; Winterscheid & Gregor, pl. 1, ig. 3. M a t e r i a l. SMNS P 1952/50, 186, 334+335 (part and counterpart); NMA 2006-105/1497, 2006-106/1497. D e s c r i p t i o n. Narrow linear leaves, l × w = 85–92 (? 84–100) × 6–9 mm, l/w ratio = ? 9–16, symmetric, base decurrent, petiole not clearly delimited from base, at least 20 mm long, straight; apex in all specimens incomplete, margin simple serrate, wavy due to its rather widely spaced, small and acute teeth, or almost entire; midrib straight to gently bent. M i c r o m o r p h o l o g y. Cuticles of both sides thin, abaxially even more delicate than adaxially, preserved in tiny fragments; anticlines of non-modiied epidermal cells straight to somewhat bent, abaxially more slender than adaxially, non-modiied cells 18–36 µm across, Unangemeldet Heruntergeladen am | 10.04.17 15:37 349 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 usually four-celled, rarely two- or three-celled prominent trichome bases, inner diameter (i.e., without margin) 13–24 µm, ± distinctly surrounded by a ring of well-cutinised, thickwalled small cells (sometimes even the cells in the wider surrounding area of a trichome base appear more strongly cutinised); stomata only exceptionally preserved and vaguely discernable, anomocytic nearly roundish, 20–22 µm long (n = 2), aperture a short, narrow oval. R e m a r k s. Specimen SMNS P 1952/50 was found determined erronously as Sideroxylon. The successfully prepared tiny cuticle fragments mostly yield a single trichome base and the immediately surrounding area. Judging from these fragments and the cell pattern relected on them, the area surrounding the trichome base is more strongly cutinised than the remaining cuticle. Diagnostically relevant cuticular features rarely have been described (Mai & Walther 1978, Kvaček 2004a). In the material from Haselbach (Mai & Walther 1978), the trichome bases are two-celled. In Flörsheim, Kvaček (2004a) notes two- to mainly four-celled trichome bases, which matches well with the here-described material in which four- and three-celled bases are more common than two-celled ones. Myrica lignitum (Unger) Saporta differs by having clearly two-celled trichome bases. In the Eocene to early Oligocene, M. longifolia was widespread from Central Europe to the Tethys and Mediterranean region, including Geiseltal, Häring, Socka, and Armissan, and is often interpreted as indicative of drier climatic conditions (Rüfle 1976). Myrica obliquifolia Kovar-Eder sp. nov. Pl. 4, igs 12–17; Pl. 15, igs 3–9 ? 2004a Myrica cf. joannis Ettingshausen 1858 emend. Kovar-Eder 1996; Kvaček, p. 11, pl. 9 igs 9, 10, 18, 19. Holotype 1952/85. designated h e r e. SMNS P P a r a t y p e s d e s i g n a t e d h e r e. SMNS P 1952/162, 232, 238, 255; SMNS P 1953/99. F u r t h e r m a t e r i a l. SMNS P 1952/63+99 (part and counterpart), 204, 386. NMA 2006142/1497. D e r i v a t i o n o m i n i s. Referring to the somewhat asymmetric leaf blade. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. D i a g n o s i s. Ovate, oblong to elliptic leaves, lamina somewhat asymmetric, margin entire near base, towards apex simple serrate with widely spaced, minute teeth; two-celled trichome bases narrow oval to 8-shaped, joint anticline of base cells very delicate. D e s c r i p t i o n. Leaves ovate, oblong to elliptic, typically somewhat asymmetric, texture thin to medium thick, l × w = 62 (? 65)–110 (? 120) × 10–25 mm, l/w ratio = 4.1– about 7, base angle acute, base straight, cuneate, petiole straight, up to 18 mm long (SMNS P 1952/85), apex long attenuate, apex angle acute, margin basally entire, then minutely simple serrate, teeth widely spaced, density higher apically, teeth narrow, apex acute, sinus acute or rounded; midvein straight or somewhat bent, secondaries craspedodromous or semicraspedodromous (SMNS P 1952/63+99), arising at about 60° angle (SMNS P 1952/85). M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle delicate, anticlines of non-modiied cells straight to somewhat curved, non-modiied epidermal cells adaxially 18–36 (average 24–25) µm, abaxially 16–45 (average 24–28) µm across, two-celled trichome bases rare, oval to 8-shaped, margin strongly cutinised, inner diameter 14–19 µm, joint wall of base cells very delicate, peltate trichomes rarely attached, 75–105 µm in diameter (n = 3), margin minimally undulate; abaxial cuticle: stomatal complexes anomocytic, stomata elliptic, l × w 20–26 × 17–27 µm, aperture elliptic to roundish, 6–8 µm long, poles acute. R e m a r k s. Specimens SMNS P 1952/85, 162, 232, and 255 are the most complete ones. Except for specimen SMNS P 1952/162, which has been transferred to artiicial resin, cuticle details are available. The micromorphology (stomata and peltate trichomes attached to two-celled bases) is suitable for generic identiication. These leaves differ from the hitherto known fossil species of Myrica by their somewhat asymmetric leaf shape combined with minute and widely spaced teeth. The anticline separating the two Unangemeldet Heruntergeladen am | 10.04.17 15:37 350 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 base cells is remarkably delicate, unlike that of, for example, Myrica lignitum or M. undulatissima (Knobloch et Kvaček 1976). Myrica cf. joannis from Flörsheim (Kvaček 2004a) differs from M. joannis Ettingshausen 1858 emend. Kovar-Eder 1996 (Kovar-Eder 1996) by having more abundant and bigger teeth. Moreover, M. joannis bears variably straight to undulate anticlines of the non-modiied epidermal cells both adaxially and abaxially. In contrast, M. obliquifolia shows straight to curved anticlines. It cannot be ruled out, however, that we are dealing here with sun leaves and that the more delicate shade leaves with possibly more undulate anticlines are not preserved in Rauenberg. In this context, note that Myrica cf. joannis from Flörsheim shows undulate anticlines. Peltate trichomes appear to be rare in both M. joannis and M. obliquifolia. An asymmetric base is described for Myrica hakeaefolia (Unger) Saporta sensu Engelhardt from Markvartice (Bůžek et al. 1976). These specimens are very fragmentary, the cuticle is not available, and the authors hesitate about the generic afinity. Juglandaceae A. Rich ex Kunth Carya Nutt. Carya fragiliformis (Sternberg) Kvaček & Walther Pl. 6, ig. 22 1825 Phyllites fragiliformis, Sternberg, p. 42, pl. 50, ig. 1. 2004a Carya sp.; Kvaček, p. 10, pl. 9 ig. 1. 2007 Carya fragiliformis (Sternberg) Kvaček & Walther comb. nov.; Walther & Kvaček, p. 110, pl. 11, igs 1–3, pl. 23, igs 8–10. M a t e r i a l. SMNS P 1952/58. D e s c r i p t i o n. Single, fragmentary, probably sessile lealet, texture thin, laminar shape asymmetric, ? oblong to minimally obovate, l × w = 70 (? 100) × 30 (? 35) mm, l/w ratio about 2.8, base asymmetric, base angle wide acute, base weakly convex, apex not preserved; margin sharply serrate, teeth with acute sinus and apex, basal and apical sides variable, near base straight, convex or concave, and in higher parts of the lamina lexuous to retrolexuous; midvein straight, secondaries densely spaced, craspedodromous to semicraspedodromous, arising at wide angles of about 80° near the base, higher up angle of origin decreasing, secondaries smoothly bent, subparallel towards each other, sending abaxially further veinlets into the teeth below, terminating in marginal teeth, tertiaries percurrent, course almost straight to somewhat sinuous, angle obtuse. R e m a r k s. This remain is a faint leaf impression with well-preserved venation. It is one of the rare leaf records of temperate deciduous taxa from Rauenberg. The differentiation of Carya fragiliformis and C. serrifolia is still open (Walther & Kvaček 2007). Carya quadrangula (Kirchheimer) Leroy Pl. 6, ig. 21 2004a Carya quadrangula (Kirchheimer) Kvaček, p. 11, pl. 9, ig. 2. Leroy; M a t e r i a l. SMNS P 1952/159, SMNS P 1953/102. D e s c r i p t i o n. Somewhat lattened endocarps, elliptic-ovoid in shape, l × w = 36 × 25 mm, apex conical-acuminate, base rounded to weakly truncate, surface smooth; specimen SMNS P 1952/159 with two narrow, long lacunae and the separating septum. R e m a r k s. The two specimens are not only very similar in shape and size to each other but are also strikingly similar to the specimen described from Flörsheim (Kvaček 2004a). Whether Carya ventricosa (Schlotheim) Schimper and C. cf. rostrata (Schlotheim) Schimper reported from the Alzey Formation (Mallison 2002) are conspeciic with the here-described specimens remains open. Engelhardia Lesch. Ex Blume Engelhardia orsbergensis (Wessel & Weber) Jähnichen, Mai & Walther Pl. 6, igs 15–17 1855 Banksia orsbergensis Wessel & Weber, p. 146, pl. 25, igs 9a–d. 1977 Engelhardia orsbergensis (Wess. & Web.) Jähnichen, Mai & Walther comb. nov., p. 326–341. M a t e r i a l. SMNS P 2228/1. SMNK 7499, 7609+7610 (part and counterpart). NMA 2006108/1497. Unangemeldet Heruntergeladen am | 10.04.17 15:37 351 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 D e s c r i p t i o n. Lealets, laminar shape oblong, l × w = 38–70 × 7–12 mm, base convex to shortly cuneate, somewhat asymmetric, no petiole, apex angle acute, apex straight, margin simple serrate, teeth distant, tiny, acuminate, midvein straight to gently bent, secondaries and ? intersecondaries arising at >45° angles and 1–3 mm distances (NMA 2006-108/1497). R e m a r k s. Though no cuticle is available, these are unambiguous records of Engelhardia orsbergensis lealets. The assignment is based on the asymmetric base and the absence of a petiolule, which makes it probable that we are dealing with lealets that were directly attached to the rhachis. The leaf base variability is well within that of E. orsbergensis, as has been demonstrated by Jähnichen et al. (1977) in their comprehensive study of Englhardia from the Palaeogene and Neogene of Europe. Some more leaf fragments may be assigned to either Engelhardia or Myrica. Engelhardia macroptera (Brongniart) Unger Pl. 6, ig. 18 1828 Carpinus macroptera Brongniart, p. 48, pl. 3 ig. 6. 1866 Engelhardia macroptera (Brongniart) Unger, p. 52, pl. 16, igs 9–11. M a t e r i a l. SMNS P 1952/107, 154+155 (part and counterpart), 206. D e s c r i p t i o n. Incompletely preserved trilobate involucres, lobes entire-margined, narrow, apex rounded, medial lobe 22–33 mm long, lateral one 18 mm long (specimen SMNS P 1952/154), medial lobe somewhat obovate, 6 mm wide in its widest part, width of lateral lobes up to 4 mm, at the base of the involucres only the position of the nutlets is visible; venation (SMNS P 1952/154): medial lobe with three veins running parallel to the lobe length, central vein stronger than lateral ones, further venation reticulate, lateral lobes also with at least two main veins and reticulate further venation. R e m a r k s. Though rare and mainly fragmentarily preserved, these involucres add evidence for the presence of Engelhardia in this assemblage. This is remarkable because neither a leaf nor a fruit have been reported from the lora of Flörsheim (Kvaček 2004a). ? Rosales ? Rhamnaceae Juss. ? Berchemia Necker ex de Candolle ? Berchemia altorhenana Kovar-Eder sp. nov. Pl. 5, ig. 5; Pl. 16, igs 7–9 Holotype 1953/92. designated h e r e. SMNS P S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. Referring to the geographical position of the fossil site in the Upper Rhine Graben. S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. D i a g n o s i s. Petiolate leaf, small, ellipticoblong, base convex, apex rounded, entiremargined to faintly crenate, secondary veins delicate, numerous, regularly spaced; anticlines deeply undulate, undulation rounded to zig-zag, stomatal complexes probably anomocytic, stomata narrow oval, marginally overlapped by a cuticular ledge, front cavity narrow elliptic to narrow spindle-shaped with distinct ledges, stoma orientation probably subparallel, trichome bases with wellstaining foot cell, surrounding cells radially arranged. D e s c r i p t i o n. Leaf petiolate, petiole very slender, 6 mm long; almost complete, slightly bent near petiole base; lamina moderately coriaceous, l × w = 25 × 12 mm, l/w ratio = 2.1, shape elliptic-oblong, base convex, base angle obtuse, apex rounded, uppermost apex not preserved, apex angle obtuse, margin entire to faintly crenate; midvein straight, ribbed, secondaries slender, hardly tapering towards leaf margin, densely spaced, about 2–3 mm apart, vein spacing and angle rather regular, veins running strongly parallel to each other, slightly diverging towards the margin, further details not available. Unangemeldet Heruntergeladen am | 10.04.17 15:37 352 M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle delicate, anticlines of non-modiied cells slender, smooth, deeply undulate, undulation rounded to zig-zag, non-modiied cells 24–38 (average 35) µm across, simple trichome bases scattered on both surfaces, abaxially more abundant, with foot cell surrounded by radially arranged cells, trichome foot cell 9–14 µm across, well staining; abaxial cuticle: type of stomatal complexes probably anomocytic, stomata narrow oval, marginally slightly overlapped by surrounding cuticle forming a cuticular ledge, stoma l × w = 17–25 (average 21) × 10–21 (average 15) µm, front cavity narrow elliptic to narrow spindle-shaped with distinct ledges, 10–17 µm long, stomata orientation probably partly subparallel. R e m a r k s. The dense and parallel spacing of the secondaries, which hardly taper along their course, are reminiscent of Rhamnaceae (Berchemia). Berchemia multinervis (A. Braun) Heer, known from the Miocene (Bůžek 1971), differs by its bigger leaf size. Berchemia has also been described from the Miocene of Kreuzau (Ferguson 1971), including small leaves such as our specimen. Camptodromites sp. Kvaček & Teodoridis (2011), originally described by Ettingshausen (1868) as Berchemia multinervis from Kučlín, resembles the heredescribed specimen by its dense secondary venation, but its leaves are subsessile. Unfortunately, SMNS P 1953/92 does not provide any information about the tertiary venation, which is characteristic in Berchemia, and the records mentioned in this context do not give information on cuticle structure. Cuticles of 15 southeast Asian species of Berchemia have been studied for comparison (Appendix 1). There is no match of the fossil with a single modern species, but the general cuticle characteristics may point towards afinity to Berchemia. B. afinis Hassk. is similar regarding the zig-zag course of the anticlines and the trichome bases (with a dark-staining centre and girdling cells, pl. 23, ig. 9). The stoma shape of our fossil with somewhat overlapping surrounding cuticle resembles B. lineata (L.) DC, for example, and to a lesser extent also B. kulingensis Schneider, B. philippinensis Vid., and B. racemosa S. & Z., whose stomata are deeply sunken among strongly papillaelike domed non-modiied cells (pl. 23, igs 7, 8). Regarding stoma orientation, B. loribunda J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 (Wall.) Brongn., B. lineata, B. longeracemosa Okuyama, and B. sinica Schneid. show subparallel stoma orientation resembling B. altorhenana. The abaxial cuticle surface of the latter taxa is, in contrast to our fossil, distinctly granulate or granulate-rugulate in B. lineata. The stomata of the studied modern species are smaller than in B. altorhenana, accounting for some ambiguity of the generic assignment. ? Fabales ? Fabaceae Lindl. ? Leguminocarpon Goeppert ? Leguminocarpon sp. Pl. 6, ig. 5 M a t e r i a l. NMA 2006-173/1497. D e s c r i p t i o n. Two shortly petiolate pods in opposite position attached to a fragmentarily preserved petiole; pod shape elliptic, one capsule rather complete, l × w about 20 × 6 mm, base acute, apex probably shortly acute or somewhat rounded. R e m a r k s. The systematic afinity of these pods remains ambiguous. Oxidales Elaeocarpaceae Juss. Sloanea L. Sloanea artocarpites (Ettingshausen) Kvaček et Hably Pl. 5, ig. 4 1869 Quercus artocarpites, Ettingshausen, p. 63, pl. 55, igs 19, 19a. 1976 Dicotylophyllum sparsidentatum sp. n.; Bůžek et al., p. 105, pl. X, igs 1–7, pl. XX igs 5, 6, pl. XXI, igs 1–6, text-ig. 8. 2001 Sloanea artocarpites (Ettingshausen) Kvaček et Hably comb. nov.; Kvaček et al., p. 117, pl. 4 igs 8, 9, pl. 6 ig. 8. 2004a Sloanea artocarpites (Ettingshausen) Kvaček & Hably; Kvaček, p. 15, pl. 11, igs 6–9, 13–15. M a t e r i a l. SMNS P 1952/61. D e s c r i p t i o n. Middle part of a delicate leaf, base and apex lacking, shape elliptic or ovate, somewhat asymmetric, l × w = ? 70 × 54 mm, Unangemeldet Heruntergeladen am | 10.04.17 15:37 353 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 margin wavy to crenate and minutely serrate, teeth narrow, single, rather widely spaced, apex and sinus acute; midvein slender, straight, secondaries craspedodromous to weakly brochidodromous, relatively widely spaced (up to 18 mm), originating alternately to suboppositely, gently bent, sending abmedial side veinlets towards the margin; no further morphological details or cuticle preserved. R e m a r k s. Based on the serration of the leaf margin and the acute teeth, this leaf resembles Sloanea artocarpites more closely than S. olmediaefolia (Kvaček et al. 2001). In Flörsheim, S. artocarpites is documented by several specimens (Kvaček 2004a). Further occurrences are Seifhennersdorf (Walther & Kvaček 2007), Holý Kluk Mts. (Radon et al. 2006), possibly also Bechlejovice (Kvaček & Walther, 2004), all dated to the early Oligocene. Sloanea olmediaefolia (Unger) Kvaček & Hably Pl. 5, igs 2, 3; Pl. 16, igs 4–6 1850 Artocarpidium olmediaefolium, Unger, p. 36, pl. 14, igs 1, 2. 2001 Sloanea elliptica (Andreánszky) Kvaček & Hably comb. nov.; Kvaček et al., p. 117, pl. 2 igs 1–5, pl. 4 igs 1–5, pl. 6 igs 1–7. 2008 Sloanea olmediaefolia (Unger) Kvaček et Hably; Hably & Kvaček, p. 140, ig. 1: 1–5. 2008 Eotrigonobalanus furcinervis (Rossmaessler 1840) Walther & Kvaček 1989; Winterscheid & Gregor, p. 5, pl. 1, ig. 4. M a t e r i a l. SMNS P 1952/340. NMA 2006114/1497. D e s c r i p t i o n. Leaves membranaceous, lamina slightly asymmetric, ovate to elliptic, l × w = 82–88 × 32 (34–35)–40 mm, l/w ratio = 2–2.6; petiole straight, 11 mm long, basally somewhat widened, abscission mark oblique (NMA 2006-114/1497); base cuneate to slightly concavo-convex, apex acuminate; margin wavy dentate, minutely toothed, teeth more closely spaced near base, more widely spaced apically, minute terminal glands probable on some teeth (NMA 2006-114/1497); midvein straight to slightly bent, ? 7–8 pairs of secondaries arising alternately to suboppositely, widely spaced, 8–15 mm (NMA 2006-114/1497), running slightly curved across the lamina, ending in tooth apices, craspedodromous, sending off abmedial branches that also end in marginal teeth; tertiaries forked percurrent with a tendency towards reticulate near the leaf margin, fourth-order veins randomly reticulate (NMA 2006-114/1497); lowermost pair of secondaries arising nearly oppositely at leaf base, ascending steeply near the margin (SMNS P 1952/340). M i c r o m o r p h o l o g y. Adaxial cuticle thick, anticlines distinctly undulate, forming puzzlelike cells 17–28 (average 23) µm across, glabrous; abaxial cuticle more delicate, anticlines of non-modiied cells, undulate, anticlines weakly cutinised, with shorter amplitude and wave length than adaxially, 12–27 (average 17) µm across (NMA 2006-114/1497), stomatal complexes anomo- to weakly cyclocytic, subsidiary cells somewhat more strongly cutinised (more intensively staining) than the other cells (SMNS P 1952/340), stomata roundish, 11–16 (average 13) × 10–15 (average 12) µm (SMNS P 1952/340) and 9–12 (average 10) × 8–10 (average 9) µm (NMA 2006-114/1497), aperture elliptic to roundish, 3–5 (average 4) µm (in both specimens) long, simple trichome bases only above veins, rare. R e m a r k s. Leaf NMA 2006-114/1497 is among the best preserved of all the investigated material. It is not only complete but shows venation details and minute dentation. Its cuticle preservation is less favourable, probably because it was transfered to artiicial resin. Although the leaf margin of specimen SMNS P 1952/340 is not preserved, the leaf shape and secondaries are already reminiscent of Sloanea, which is conirmed by the wellpreserved cuticle. Kvaček (2004a) stressed the differences in the undulation of the anticlines abaxially and the type of stomatal complexes between Sloanea elliptica (now S. olmediaefolia) and S. artocarpites. Accordingly, the here-described leaves match S. olmediaefolia better due to the undulated anticlines and the stomatal complexes, which are cyclocytic, though weakly. The cyclocytic type is indicated by more intensively staining subsidiaries in specimen SMNS P 1952/340. The leaf shape and size variability of S. olmediaefolia from the Tard Clay between Óbuda (Budapest) and Eger-Kiseged is considerable (Hably & Erdei 2015), and our specimens are closer in size to those from Eger-Kiseged. S. olmediaefolia was known previously only from the early Oligocene of the Paratethys Province (Hungary and Slovenia) (Kvaček et al. 2001). Our record extends Unangemeldet Heruntergeladen am | 10.04.17 15:37 354 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 its geographical range during the early Oligocene considerably westwards and northwards and is a further substantiation of the loristic relationship between the early Oligocene lora of the Budapest region and that of the western coastal regions of Palaeo-Europe. are unknown for P. leuce. The relatively small size of the lamina accounts for the systematic assignment to P. germanica. Malphigiales Craigia W.W. SM. et W.E. Evans Salicaceae Mirb. Populus L. Malvales Malvaceae Juss. Craigia bronnii (Unger) Kvaček, Bůžek et Manchester Pl. 6, ig. 2 Populus germanica (Menzel) Walther Pl. 5, ig. 6 1926 Menispermites germanicus Menzel, p. 32, igs 1–3. 1978 Populus germanica (Menzel) comb. nov.; Walther, p. 90, pl. 3, ig. 16, pl. 8, igs 1–9, pl. 9, igs 1, 2, pl. 36, igs 1–8, pl. 37, igs 1–5. 2004a Populus germanica (Menzel) Walther; Kvaček, p. 15, pl. 11, igs 10–12. M a t e r i a l. SMNK 7568, ? 7725. NMA 2006107/1497. D e s c r i p t i o n. Membranaceous leaves, petiole lacking, l × w = ? 55–70 × 60–70 mm (SMNK 7568, NMA 2006-107/1497) and 9 × 12 mm (SMNK 7725), shape suborbicular, base rounded (SMNK 7568) and ? concavo-convex (SMNK 7725), apex ? rounded to obtuse, ? acuminate in the uppermost part (SMNK 7725), margin wavy (SMNK 7568); midrib straight, secondaries craspedodromous, basal secondaries arising directly at base, position of further secondaries subopposite, almost straight (SMNK 7568) or slightly bent (SMNK 7725), single secondaries forking or sending abmedial veinlets, terminating in glandular marginal teeth or lobe sinus; tertiaries percurrent, fourth-order venation reticulate, imbrial vein ± distinct along leaf margin (SMNK 7568). R e m a r k s. Specimens SMNK 7568 and NMA 2006-107/1497 are embedded in synthetic resin. This enables the leaf venation to be traced in transmitted light and reveals the existence of a imbrial vein (SMNK 7568). Because the very small lamina, the assignment of specimen SMNK 7725 may be ambiguous. Kvaček (2004a) pointed to the similarity of Populus germanica and P. leuce (Rossmässler) Unger from the Staré Sedlo Formation (Knobloch et al. 1996), stating that the two species may be conspeciic but that the cuticle structures 1845 Ulmus bronnii Unger, p. 79 pro parte, pl. 25, igs 2–4. 1991 Craigia bronnii (Unger); Kvaček, Bůžek et Manchester, p. 552. M a t e r i a l. SMNK 7748 (part and counterpart). D e s c r i p t i o n. Flattened, detached, oval valves of a fruit capsule, 12 mm long with peripheral, incompletely preserved wing, diameter 27 (? 30) mm, partly preserved as compression and partly as imprint, central suture well visible, wing poorly preserved, only main veins discernable, spreading radially, sometimes dichotomising towards margin. R e m a r k s. In this poorly preserved, rather large specimen, seeds are not demarcated and it remains open whether it was sterile. Craigia is best known from the North Bohemian brown coal region, where it is preserved in the form of different organs: fruits, foliage, lowers, and lower buds (Kvaček 2004c). This is the only record of Craigia in the lora from Rauenberg. Saxifragales Hamamelidaceae R. Br. In C. Abel Distylium Siebold et Zucc. Distylium metzleri Kovar-Eder sp. nov. Pl. 6, ig. 23; Pl. 16, igs 1–3 Holotype 1952/43. designated h e r e. SMNS P S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. Unangemeldet Heruntergeladen am | 10.04.17 15:37 355 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 D e r i v a t i o n o m i n i s. In honour of Rudolf Metzler, one of the major collectors of the lora from Rauenberg. D i a g n o s i s. Slender, smooth concentric epicuticular wrinkles developed upon guard cells and subsidiary cells, wrinkles thicker, distinct and parallel upon veins, trichome bases scattered above veins, cuticular wrinkles near trichome bases ± radially oriented. D e s c r i p t i o n. Apical two thirds of a simple, asymmetric leaf, texture thin, laminar shape elliptic or ovate, l × w = 51 (? 65) × 22 mm, apex almost straight to somewhat acuminate, apex angle acute, leaf margin entire in the central part of the leaf but sparsely and minutely denticulate near apex; secondaries faintly visible, brochidodromous, widely spaced (distance between two adjacent secondaries 13 mm), veins diverging from the secondaries towards the leaf margin, forming further loops, a single intersecondary and few tertiaries visible, the latter percurrent. M i c r o m o r p h o l o g y. Cuticles of both sides very thick and well preserved; anticlines of non-modiied epidermal cells thick, undulate, slightly knobby; undulation shallower adaxially than abaxially, non-modiied cells almost equal in size on both sides, adaxially 26–31 (average 32) µm, abaxially 23–40 (average 31) µm; abaxial cuticle: stomatal complexes brachyparacytic, somewhat asymmetric due to subsidiary cells that are partly unequal in size; pair of guard cells elliptic oblong, 18–26 (average 21) × 19–23 (average 22) µm, l/w ratio = 0.8–1.3 (average 0.9), outer front cavity narrow oval, almost reaching the poles, formed by distinct cuticular ledges, epidermal wall of guard cell not very distinct, poles somewhat thickened; upon guard and subsidiary cells, slender concentric cuticular wrinkles developed; single, big and prominent, thickly cutinised trichome bases above veins, one to two rings of small cells surrounding the trichome base centre, inner diameter about 20 µm, diameter including girdling cells about 70 µm, epicuticular wrinkles distinct, thick and radially arranged around trichome bases but running parallel above veins. R e m a r k s. This is one of the rare leaf indings in this lora showing secondary veins and tertiaries. Unfortunately, the basal part of the lamina is not preserved. The combination of features – asymmetric lamina, brachyparacytic stomata, and the characteristic trichome bases – indicate afinity to the Hamamelidaceae. Distylium fergusonii differs by its entire margin also near the leaf apex, by the abaxial cuticle which is less strongly cutinised than the adaxial one, and by the greater abundance of trichome bases. The epicuticular structure is clearly highly variable, from almost absent (Mai & Walther 1978, there as D. cf. fergusonii; Ferguson 1971) to well developed in the material from Wackersdorf (Knobloch & Kvaček 1976) and Oberdorf (Kovar-Eder & al. 2001). D. fergusonii plexus from the Kristina mine, Hrádek, bears thick cuticles on both sides, and the strong wrinkles abaxially obscure the anticlines of the non-modiied cells (Holý et al. 2012). Leaves of D. heinickei (Mai & Walther 1991) bear tiny marginal teeth, as does the here-described specimen. Studies of the type and original material revealed that the cuticle is very delicate and the epicuticular structure is less strongly developed, and in cf. Distylium sp. (Walther 1999) the cuticle is smooth. No delicate concentric crinkles have been reported on the guard and subsidiary cells in D. fergusonii or in D. heinikei. Since leaves of Distylium are rather rare in the fossil record and usually not abundant at single sites, it is dificult to judge fossil species variability. Especially for D. fergusonii the variability of the described records is broad, raising doubts about whether they all are conspeciic. ? Santalales ? Loranthaceae Juss. ? Viscophyllum Knoll ? Viscophyllum hendriksiae Kovar-Eder sp. nov. Pl. 7, igs 1–5; Pl. 23, igs 1–6 Holotype 1953/70. designated h e r e. SMNS P P a r a t y p e s. SMNS P 1952/11, 57, 158; SMNS P 1953/71, 77. NMA 2006-134/1497. S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. Unangemeldet Heruntergeladen am | 10.04.17 15:37 356 L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. In honour of Janine Hendriks, technician at NMA, for her assistence in my study of the collection there. D e s c r i p t i o n. Long-petiolate, leathery, entire-margined leaves, petiole up to at least 25 mm long and up to 3 mm thick, broader near petiole base, often bent; lamina elliptic to broad elliptic, l × w = 35 (? 38)–56 × 15–31 mm, l/w ratio = 1.6–3.4, base cuneate, gradually tapering into the petiole, apex angle acute, shape (almost) straight, utmost apex blunt; midvein straight, visibly continuing into the petiole, midvein thick and pronounced in specimen SMNS P 1952/11 but slender and hardly visible in other specimens, secondaries very delicate, widely spaced, steeply ascending and arching about 1–2 mm in front of the margin (brochidodromous) (NMA 2006-134/1497), further details unclear. M i c r o m o r p h o l o g y. Cuticle thick, glabrous, anticlines mainly straight, thick but often not well traceable, hypostomatic; adaxial cuticle: non-modiied epidermal cells polygonal, very large, up to 114 µm across, cell surface subdivided into smaller ields indicating ? hypodermis; abaxial cuticle: non-modiied epidermal cells ± distinctly arranged in rows, subquadrangular to pentagonal, with central thickened papilla, cells about 22–43 µm across (SMNS P 1952/11), that is, much smaller than stomata, or non-modiied cells polygonal as in non-stomata bearing condition, stomata ± completely surrounded by a thick cuticular fold, thus type of stomatal complexes ambiguous, stomata arranged parallelly (SMNS P 1952/11) or randomly (SMNS P 1952/158), oval to almost roundish, l × w = 43–83 × 33–57 µm, epidermal wall of guard cell indistinct, cuticular ledges very thick, forming an elliptic to broad spindleshaped, very prominent, 34–56 µm long front cavity, aperture short, narrow oval to spindleshaped, parallel epicuticular striation present, on guard cells concentric; SEM study did not yield further details. R e m a r k s. The texture of specimens SMNS P 1952/11 and 57 is extremely leathery; in SMNS P1952/158 and SMNS P 1953/71 it is less leathery but still thick. In all these specimens the carbonised leaf material is fragmented into tiny splinters, while the lamina J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 of SMNS P 1953/70 and 77 appears skin-like, unfragmented and brownish. In all specimens the cuticle is thick but mainly present in small fragments. The cuticular features leave no doubt that these specimens are conspeciic. In particular, this concerns stoma shape and size as well as the peculiar cell pattern in nonstoma bearing condition (large polygonal cells with a network of smaller meshes indicative of a ? hypodermis). In SMNS P 1952/158 the stomata are randomly distributed, while in the other specimens the stoma orientation appears rather parallel. It remains open whether the differences in stoma orientation are related to the problem that cuticle samples were obtained mainly from near the leaf margin close to the leaf base to avoid coverage by varnish. These leaves probably are assignable to the fossil genus Viscophyllum Knoll. V. weylandii (Walther) Walther (Mai & Walther 1985), described from the Upper Eocene of the Weisselster Basin, is similar in leaf shape and in the steeply ascending secondaries; it differs by its smaller leaf size, smaller stomata (which are wider than long) and the type of stomatal complexes with clearly visible subsidiaries. Cornales Hydrangeaceae Dumortier Hydrangea L. Hydrangea microcalyx Sieber 1881 Pl. 6, ig. 4 1881 Hydrangea microcalyx, Sieber, p. 16 pro parte: igs 26, 27, 31. 2014 Hydrangea microcalyx Sieber 1880; Winterscheid & Kvaček. M a t e r i a l. SMNS P 1952/145. D e s c r i p t i o n. Tetramerous, petaloid calyx, 17 mm in diameter, single calyx leaves elliptic to roundish in shape, tapering towards base, point of attachment to axis narrow, apex obtuse or somewhat emarginate, entiremargined, every leaf shows a midvein running straight into the apex, especially the basal secondaries arise at narrow angles, ascending steeply, eventually forking across the lamina, further details not available. R e m a r k s. Only a single specimen is available from the abundant material which is Unangemeldet Heruntergeladen am | 10.04.17 15:37 357 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 surprisingly well preserved. Its four leaves are unfused, closely spaced and touching each other. The specimens from Kučlín (Sieber 1881, Kvaček & Teodoridis 2011) differ in that the leaves are obovate and not as closely spaced as in the specimen from Rauenberg. The latter more closely resembles the specimen from Seifhennersdorf igured by Mai (1963: pl. 10, ig. 9) and the specimen from Hammerunterwiesenthal (Walther 1998: Abb. 2 ig. 20). The specimens from Seifhennersdorf (Walther & Kvaček 2007: pl. 13 igs 7, 8) are incomplete and thus not well comparable. This large variability, however, is within the natural variability of Hydrangea species Mai (1963). H. microcalyx is an accessory element in the volcanic loras of Bohemia and Saxony from the late Eocene to Oligocene. The revision of the lora from Orsberg near Bonn also includes H. microcalyx (Winterscheid & Kvaček 2014). This is the irst record from the Oligocene coastal loras of the Rhein Graben. The east Asian H. paniculata Sieb. et Zucc., H. petiolaris Sieb. et Zucc., and H. quercifolia Bartr. from North America are regarded as the most similar relatives (Mai 1963). Ericales ? Pentaphyllaceae Engl. in Engl. & Prantl M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle thick, glabrous; anticlines of nonmodiied epidermal cells densely undulate but not very pronounced, cells 32–49 (average 42) µm across; abaxial cuticle: anticlines of non-modiied epidermal cells even less distinct than adaxially, undulate but faintly traceable only (cell size not measureable), type of stomatal complexes unclear, stoma elliptic, stomata surrounded by a thick cuticular wrinkle, guard cells less strongly cutinised, occasionally with concentric striae, cuticular ledges strongly thickened, forming a narrow oval outer front cavity, stoma l × w = 32–45 (average 38) × 26–35 (average 30) µm, l/w ratio = 1.1–1.5 (average 1.3), aperture length 24–35 (average 29) µm, cuticle surface with scattered, almost parallel wrinkles. R e m a r k s. Laminar shape and size remain ambiguous due to the fragmentary preservation of this leaf. The cuticle suggests a possible relationship to the Pentaphyllaceae. Stoma shape and size, the concentric wrinkles around the stomata, and the occasionally occurring, ± parallel wrinkles covering the abaxial surface may point towards afinity to Gordonia. The very dense undulation of the anticlines on the adaxial cuticle raises uncertainty, however. ? Ternstroemites Berry emend. Hickey Symplocaceae Desf. ? Ternstroemites maritiae Kovar-Eder sp. nov. Symplocos Jacq. Pl. 5, ig. 7; Pl. 17, igs 1–3 Symplocos deichmuelleri (Kvaček & Walther) comb. nov. Holotype 1952/342. designated h e r e. SMNS P S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. Dedicated to Marit Kamenz, technician at the SMNS, who patiently made numerous cuticle preparations for this study. D e s c r i p t i o n. Apical fragment of a coriaceous, probably entire-margined leaf, l × w = 35 (? 70) × 22 (? 24) mm, laminar shape unclear, apex angle acute, apex shape nearly straight. Pl. 5, igs 8–10; Pl. 17, igs 4–6 1998 Dicotylophyllum deichmuelleri Kvaček & Walther, p. 14 pl. 6, igs 7–12. 2004a Symplocos volkeri Kvaček sp. n., Senckenberg Lethaea, 84, p. 17, pl. 14, igs 2, 3, 13–15. 2007 Dicotylophyllum deichmuelleri Kvaček & Walther; Walther & Kvaček, p. 127, pl. 18, igs 16, 17, pl. 24, igs 12, 13. 2008 Eotrigonobalanus furcinervis (Rossmaessler 1840) Walther & Kvaček 1989; Winterscheid & Gregor, pl. 1 ig. 4. 2014 Symplocos volkeri Kvaček 2004; Winterscheid & Kvaček, p. 30, pl. 5 igs 7, 8, pl. 12 ig. 1. 2014 Dicotylophyllum deichmuelleri Kvaček et Walther; Kvaček et al., p. 42, igs 7G–J, 8L. M a t e r i a l. SMNS P 1952/62, 65, 79+98 (part and counterpart), SMNS P 1953/43. Unangemeldet Heruntergeladen am | 10.04.17 15:37 358 D e s c r i p t i o n. Petiolate leaves, texture medium thick, l × w = 40–55 (? 58) × 17–33 mm, l/w ratio = 1.8–2.8, laminar shape broad elliptic to slightly obovate, petiole 4–9 (? complete) mm long, bent or straight, basally widened (SMNS P 1952/65); base angle acute to almost 90°, base rather straight to concave near the very base and then somewhat convex, apex angle acute to obtuse, apex straight to somewhat acuminate; margin almost entire near the base with only single teeth, simple serrate with more densely spaced teeth towards apex, teeth ± tiny, basal and apical sides almost straight, sinus rounded or acute, apex rounded or (bluntly) acute and glanduliferous; midvein straight, secondaries markedly more delicate than midvein (SMNS P 1952/62), arising at about 40–50° angle from midvein, angles of adjacent veins variable, near their origin running almost parallel to midvein before turning outwards, tertiaries percurrent, course slightly convex to sinuous, higher-order veins forming a dense network. M i c r o m o r p h o l o g y. Adaxial cuticle delicate, abaxial cuticle thick, both sides glabrous; adaxial cuticle with faint anticlines, straight to bent, non-modiied cells 18–33 µm across; abaxial cuticle: non-modiied cells distinct, smaller than stomata, 13–26 (average 17–18) µm across, surface of non-modiied cells papilla-like thickened, anticlines straight to curved, stomatal complexes brachyparacytic, stomata not sunken, narrow oval, only faintly if at all delimited from narrow subsidiaries, stoma l × w = 19–29 (average 21–24) × 14–18 µm, aperture narrow oval, 6–9 µm long, cuticular ledges distinct, almost reaching the poles, slender polar I-pieces sometimes present, faint concentric striation on stomata (SMNS P 1952/62). R e m a r k s. Specimens SMNS P 1952/62 and SMNS P 1953/43 show secondary veins near the base. In the latter the tertiary and higherorder venation is also well visible, while its cuticle is less well preserved than in the other remains. The cuticle is best preserved in specimen SMNS P 1952/65. In SMNS P 1952/62 the marginal teeth in the apical part of the lamina are bigger and the apical glands on the teeth are distinctive. In the other specimens the glanduliferous teeth are easy to miss. The features of the abaxial cuticle of Symplocos volkeri and Dicotylophyllum deichmuelleri J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 are very similar and Kvaček et al. (2014) already suggested Symplocos as the probable genus for the leaves assigned to D. deichmuelleri. Comparing the leaves from Flörsheim and Rott (Kvaček 2004a, Winterscheid & Kvaček 2014), S. deichmuelleri shows considerable variability in leaf shape and secondary venation but consistency regarding the cuticular structures. The igure of Kvaček (2004a: pl. 14, ig. 14) suggests papilla-like thickenings on the surface of non-modiied cells, as in the material from Rauenberg. This was conirmed by comparative studies of the original cuticular slides from Flörsheim. Although smaller than the leaves from Flörsheim, those from Rauenberg more closely resemble the record from Flörsheim than the specimens from Rott. No glands have been reported from the tooth apices of specimens from Flörsheim or Rott. More material is therefore necessary to decide whether we are dealing with a single species. Gentianales Apocynaceae Juss. Trachelospermum Kvaček Trachelospermum kelleri Kovar-Eder sp. nov. Pl. 6, igs 24, 25; Pl. 18, igs 3–5 H o l o t y p e. NMA 2006-121/1497. P a r a t y p e. SMNS P 1953/101. S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. In honour of the private collector Manfred Keller, who donated his collection to the Naturmuseum Augsburg. D i a g n o s i s. Long-petiolate, narrow oblong to slightly obovate, entire-margined leaves, base acute to cuneate, apex acute, venation delicate; cuticles delicate, small mesophyllous secretory bodies very dense, remains of ? guard cells and secretory material masking the cuticular features of the stomata. Unangemeldet Heruntergeladen am | 10.04.17 15:37 359 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 D e s c r i p t i o n. Long-petiolate, entire-margined, coriaceous leaves; petiole 11–14 mm long (almost complete), straight to slightly bent, basally slightly widened and with longitudinal folds (NMA 2006-121/1497); base cuneate, base angle acute, shape straight; apex angle acute, shape straight (NMA 2006-121/1497); lamina narrow, oblong or slightly obovate, l × w = 75 (? 80–85) × 15 mm, l/w ratio = ? 5.3– 5.7 (SMNS P 1953/101), l × w = 38 × 11 mm, l/w ratio = 3.4 (NMA 2006-121/1497); venation delicate, midvein straight, secondaries slender, connecting at different distances from margin, intersecondaries present; secondaries, intersecondaries and tertiaries forming irregular vein network with meshes of different sizes and shapes. M i c r o m o r p h o l o g y. Cuticle of both sides delicate, non-modiied cells not discernable, cuticular features of stomata characteristically masked by intensively staining remains of the ? guard cells and secretory material, stomatal complexes ? anomocytic, stomata (broadly) oval, somewhat oblique in shape, recessed at poles, l × w = 24–30 × 19–27 µm (SMNS P 1953/101, n = 4) and 21–28 (average 24) × 15–18 (average 17) µm (NMA 2006-121/1497), length of stomatal aperture 17 µm (n = 1); densely packed with mesophyllous, differently sized secretory bodies up to 12 µm in diameter (NMA 2006121/1497). R e m a r k s. These remains are to be assigned to the genus Trachelospermum as deined by Kvaček (2004a). They differ in leaf shape and venation from T. steiningeri. Micromorphological differences are the absence of anticlines of non-modiied cells, the presence of ? guard cell remains and secretory material masking the stomatal cuticular features, the shape of the stomatal aperture, and probably also the type of stomatal complex. Apocynaceae gen. (Kvaček 2004a) is similar in leaf shape, the presence of dense secretory bodies in the mesophyll, and stoma shape. It differs from the here-described specimens by the short petiole and dense spacing of the secondaries. Although Apocynaceae gen. shows similarity to Dicotylophyllum sp. 11 from Kundratice (Kvaček & Walther 1998) in leaf shape and venation, the cuticles of the latter species differ by the type of the stomatal complex, aperture shape, idiocuticular striation and the lack of mesophyllous secretory bodies. Trachelospermum steiningeri Kvaček Pl. 6, igs 26, 27; Pl. 17, igs 7–9; Pl. 18, igs 1, 2 2004a Trachelospermum steiningeri, Kvaček, p. 18, pl. 15, igs 1–8, 11–15. M a t e r i a l. SMNS P 1952/101, 366. D e s c r i p t i o n. One complete leaf (SMNS P 1952/101) and one fragment lacking base, apex and margin (SMNS P 1952/366), moderately coriaceous; SMNS P 1952/101: laminar shape wide elliptic, l × w = 38 × 25 mm, l/w ratio = 1.5, petiole straight, 8 mm long, incomplete; base concave, base angle acute, apex angle obtuse, apex retuse, margin entire, somewhat undulate; midvein straight, distinctly tapering apically, secondaries slender, arising at angles of about 50–60°, angles decreasing apically, distance between secondaries up to 5 mm (in the central part of the lamina); course slightly diverging between each other and somewhat concave near the margin, intersecondaries probably present. M i c r o m o r p h o l o g y. SMNS P 1952/101: cuticle of both sides delicate, glabrous, anticlines only faint, smoothly curved to somewhat wavy, non-modiied epidermal cells 18–34 (average 25) µm across adaxially and 17–20 µm (n = 3) across abaxially, small secretory bodies very dense, diameter 10–15 µm; abaxial cuticle: type of stomatal complexes questionable, stomata broad oval to roundish, l × w = 14–18 × 12–19 µm, l/w ratio = 0.9–1.2 (n = 5), epidermal wall of guard cells hardly developed, guard cells hardly staining, stomatal ledges slender, slightly thickened, forming short, broad spindle-shaped front cavity with acute poles, length of front cavity 9–10 µm, aperture a faint slit. SMNS P 1952/366: adaxial and abaxial cuticle well preserved, glabrous, smooth, anticlines of non-modiied cells straight to moderately bent, slender, non-modiied epidermal cells polygonal, on both sides very similar in size, 19–34 (average 25) µm across; abaxial cuticle: stomatal complexes cyclocytic, subsidiary cells similar in size and shape to non-modiied cells but less strongly cutinised, stomata elliptic, l × w = (22) 25–31 (average 25) × 19–23 (average 21) µm, l/w ratio = 1.2–1.6 (average 1.3), epidermal wall of guard cells faintly visible only, ledges slender, not strongly cutinised, forming an elliptic front cavity, length Unangemeldet Heruntergeladen am | 10.04.17 15:37 360 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 (7) 11–17 (average 14) µm, aperture slit-like, poles I-like, slightly thickened; small, secretory bodies present. R e m a r k s. The gross morphology of specimen SMNS P 1952/101 points to afinity to Trachelospermum, but specimen SMNS P 1952/366 is too fragmentary for any assignment based on gross morphology. In both specimens the shape of the stomata and front cavity as well as the small secretory bodies, which are dense in specimen SMNS P 1952/101, support this assignment. The cyclocytic stoma complex type is evident in SMNS P 1952/366 only. Kvaček (2004a) described it as anomocytic to incompletely cyclocytic for the specimens from Flörsheim. The different interpretations are not contradictory but probably result from the fact that the cuticles from Flörsheim were unstained and that the thinly cutinised subsidiaries are probably more dificult to discern there. In specimen SMNS P 1952/101 the stomata are smaller and more roundish than in specimen SMNS P 1952/366 (therefore described separately), but in both the stoma length is within the variability provided by Kvaček (2004a). However, no giant stomata have been observed in the here-studied material. At irst glance the adaxial cuticle of SMNS P 1952/366 gives the impression of domed nonmodiied cells, but this interpretation relects the dense remains of small secretory bodies. Trachelospermum steiningeri is known so far only from Flörsheim and Rauenberg, and the number of available specimens is still very limited, leaving intraspeciic variability poorly understood. Kvaček (2004a) compares T. steiningeri to T. asiaticum (Sieb. & Zucc.) Nakai, an evergreen woody liana of upland forests and brushwood, and to further Trachelospermum species in SE Asia. Lamiales Oleaceae Hoffmanns & Link Oleinites Cookson emend. Sachse Oleinites altorhenana Kovar-Eder sp. nov. Pl. 5, igs 17, 18; Pl. 18, igs 6–9 Holotype 1952/191. designated h e r e. SMNS P P a r a t y p e s. SMNS P 1952/388; SMNS P 1953/62. S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. Derived from the geographical position of the fossil site in the Upper Rhine Graben. D i a g n o s i s. Broad elliptic to ovate, mediumsized leaves, non-modiied cells abaxially remarkably small, stomata densely spaced, subcircular, epidermal and poral wall of guard cells indistinct, prominent simple bases of probably peltate trichomes with rim-like thickened margin. D e s c r i p t i o n. Leaves elliptic to ovate, texture moderately thick, l × w up to 115 × 43 mm, l/w ratio = 2.6 (SMNS P 1953/62), margin entire to smoothly undulate, base angle obtuse, shape somewhat convex, apex angle acute, shape acuminate; midvein slender, straight, secondaries poorly visible, delicate, originating at about 50° angle, vein spacing about 10 mm in the basal half of the lamina, secondaries running almost straight close to the margin (SMNS P 1953/62). M i c r o m o r p h o l o g y. Adaxial cuticle less robust than abaxial one, unequivocally preserved only in SMNS P 1952/191; both surfaces inely striate along veins; adaxially, anticlines of non-modiied epidermal cells curved to somewhat sinuous, 17–27 (average 21) µm across, while abaxially distinctly smaller, 6–17 (average 12) µm across; in SMNS P 1952/388 anticlines of non-modiied cells not traceble; abaxial surface: stomatal complexes anomocytic, stomata more weakly cutinised than the other parts of the abaxial surface, epidermal and poral wall of guard cells indistinct, stomata roundish, l × w = 16–21 (average 18) µm × 16–23 (average 18) µm, aperture broad elliptic to roundish, 6–9 (average 8) µm long; simple trichome bases scattered, inner diameter 5–13 µm, margin collar-like, 4–9 µm thick, surrounding cells ± radially arranged, occasionally with faint radial folding, trichome heads disc-shaped or globular about 20–25 µm in diameter (? and bigger). Unangemeldet Heruntergeladen am | 10.04.17 15:37 361 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 R e m a r k s. The most complete specimen is SMNS P 1953/62 but the cuticle is best preserved in SMNS P 1952/191, which represents only the apical half of a leaf. The adaxial cuticle either had less preservation potential or else the unclear details of the cell outlines hamper its recognition. Only in specimen SMNS P 1952/191 is the adaxial cuticle unambiguous. The type of trichome base as well as the roundish, weakly cutinised and thus indistinct stomata point to the morphogenus Oleinites (Sachse 2001). This new fossil species differs from Oleinites hallbaueri (Mai) Sachse by the entire margin of the leaves, and from O. hallbaueri and O. maii (Bůžek, Holý & Kvaček) Sachse by the small, roundish stomata which are hardly delimited from the non-modiied cells with short, elliptic to roundish aperture, and by an adaxial cuticle that is usually dificult to identify. O. pachyphyllus (Kvaček 2004a), described from Flörsheim, differs both in gross morphology (elongate, narrower leaf) and in cuticular features (much thicker cuticles, larger non-modiied cells and large stomata). Oleinites altorhenana more closely resembles O. liguricus from Torre Sterpi, northern Italy, Messinian (Sachse 2001) both macroand micromorphologically. As in the hereintroduced fossil species, the adaxial cuticle of L. liguricus is dificult to discern. Apart from the difference in age, O. liguricus differs by having somewhat larger stomata. Among modern genera, similar trichome bases and roundish stomata occur in Chionanthus and Olea (Sachse 2001). Oleinites rauenbergensis Kovar-Eder sp. nov. Pl. 5, igs 12–16; Pl. 19, igs 1–6 Holotype 1952/380. designated h e r e. SMNS P P a r a t y p e s. SMNS P 1952/299, 402. F u r t h e r m a t e r i a l. ? SMNS P 1952/6; ? SMNS P 1953/93 (pl. 5, igs 15, 16). S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. Referring to the type locality. D i a g n o s i s. Small, spatulate, entire-margined coriaceous leaves, cuticle very thick, brittle, non-modiied epidermal cells very large, anticlines undulate, stomatal complexes subcircular, widely spaced, epidermal wall of guard cells faintly developed but poral wall well developed; simple tichome bases prominent. D e s c r i p t i o n. Leaves obovate, spatulate, texture coriaceous, l × w = 13–20 (? 25) × 4–11 mm, l/w ratio = 2.5–3.3, base cuneate to decurrent with gradual transition into a ? short petiole, apex rounded to somewhat emarginate, margin entire, midvein slender, straight, no further details available. M i c r o m o r p h o l o g y. Adaxial cuticle very thick, abaxial cuticle somewhat thinner, cuticle of both sides characteristically brittle, breaking into tiny fragments, simple trichome bases scattered, abaxially probably more common than adaxially, with strongly thickened, collarlike margin (up to 10 µm), surrounding cells ± radially arranged, thickened, sometimes radially elongated, a single small head found attached, 39 µm in diameter (SMNS P 1952/380); adaxial cuticle: anticlines up to 3–4 µm thick, widely and shallow undulate, non-modiied cells 22–64 (average 36–48) µm across; abaxial cuticle: anticlines of non-modiied cells poorly visible, stomatal complexes ? anomocytic, widely spaced, subcircular, epidermal wall of guard cells poorly developed, ledges thickened, forming a broad elliptic front cavity with acute poles, short, slender polar I-pieces sometimes weakly developed, stoma size l × w = 22–30 × 20–26 µm (in all specimens only few stomata available), length of front cavity 10–14 µm, double-contured. R e m a r k s. The cuticle of this fossil species is preserved mainly in very tiny fragments representing few cells, and only a few stomata were suitable for taking measurements. From the smallest leaf (SMNS P 1952/299, l × w = 13 × 4 mm) the adaxial cuticle is available, and the abaxial side probably is represented by a tiny fragment with a single trichome base. Based on the obovate, spatulate lamina along with the thick and brittle cuticle, the undulated course of the anticlines adaxially and the trichome base, this specimen is grouped here. Nonetheless, adaxially the non-modiied Unangemeldet Heruntergeladen am | 10.04.17 15:37 362 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 epidermal cells are smaller (average 36 µm). In specimen SMNS P 1952/6 it is unclear whether the trichome bases are one-celled. Specimen SMNS P 1953/93 is similar in gross morphology except for the distinctly thickened leaf margin. From this specimen only the adaxial cuticle is available. The size and pattern of the non-modiied cells correspond to those of Oleinites rauenbergensis. These spatulate small leaves are reminiscent of Ericaceae and especially of the recently described Andromediphyllum ungeri from Flörsheim (Kvaček 2004a). The cuticle, however, differs by its brittleness, much bigger cells adaxially, the thick abaxial cuticle with smaller stomata lacking distinct striae, as well as the presence of trichome bases of the Oleinites type. Oleinites dieteri (Kovar-Eder) Kvaček (Kvaček 2004a) from early Miocene lignite deposits of Styria (Kovar-Eder & Meller 2001) is similar in leaf size and shape but differs by the acute to acuminate apex and marginal serration. The cuticle of the latter species is brittle and the cells of the adaxial cuticle are similarly large with undulate anticlines, as in the here-described material. Moreover, the stomata resemble O. rauenbergensis in shape and size and peltate trichomes are present. The strong idiocuticular striation, however, distinguishes O. dieteri from O. rauenbergensis. The Oleaceae afinity of these small spatulate leaves continues to be disputable. spinose; midvein straight, secondaries craspedodromous, intersecondaries present, adjacent ones looping, secondaries and intersecondaries arising at angles of >60°, tertiaries and higherorder veins polygonal reticulate, with marginal imbrial vein. R e m a r k s. This leaf is preserved as a faint imprint with little carbonized material especially along the veins and leaf margin. The leaf surface is covered by abundant roundish fruiting bodies of fungi. Gross morphologically, Pungiphyllum waltheri from the Eocene plant site Eckfeld differs by the pattern of the tertiary venation where the tertiaries arise from the midvein between the secondaries (Frankenhäuser & Wilde 1995). The here-described specimen closely resembles the specimen from Kundratice igured by Kvaček & Walther (1981: pl. 7 ig. 1), for example, where the tertiaries originate perpendicularly mainly from the secondaries, forming a polygonate network. The single remain reported from Flörsheim (Kvaček 2004a) represents the apical half of a leaf. Although P. cruciatum is a characteristic element of Oligocene and Miocene plant assemblages, it occurs mainly as an accessory element. Dicotylophyllum incertae sedis Dicotylophyllum badense Kovar-Eder sp. nov. Pl. 7, ig. 16; Pl. 19, igs 7–9 Fam. indet. Pungiphyllum Frankenhäuser & Wilde Pungiphyllum cruciatum (Al. Braun) Frankenhäuser & Wilde Pl. 5, ig. 11 Pungiphyllum cruciatum (Al. Braun 1851) nov. comb.; Frankenhäuser & Wilde, p. 101. 2004a Pungiphyllum cruciatum (Al. Braun) Frankenhäuser & Wilde; Kvaček, p. 19, pl. 16, ig. 11. 1995 M a t e r i a l. NMA 2006-126/1497. D e s c r i p t i o n. Basal half of lamina, length and width incomplete, l × w = 55 × 25 mm, base cuneate, base angle acute, base almost straight, petiole 5 mm long, incomplete, straight, laminar shape not evident, with deep lobation, two lobes preserved, sinus widely rounded, apex Holotype 1952/200. designated h e r e. SMNS P S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. T y p e l o c a l i t y. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. Rauenberg, the fossil site, is located in Baden, Baden-Württemberg. D i a g n o s i s. Leaf coriaceous, narrow, entiremargined, cuticles thick, anticlines straight to curved, stomatal complexes cyclocytic, oval to roundish, front cavity formed by distinct cuticular ledges, reaching the poles, polar I-thickenings moderately developed, aperture short oval, simple trichome bases rare. Unangemeldet Heruntergeladen am | 10.04.17 15:37 363 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 D e s c r i p t i o n. Fragment of the middle part of a narrow leaf lacking base and apex, l × w = ? 40 × 18 mm, entire-margined, midvein somewhat bent, texture very coriaceous. M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle thick, anticlines straight to curved, shape and size of non-modiied cells on both sides very similar, 15–30 µm (average 23) µm across; simple, well-cutinised trichome bases abaxially and ? adaxially present but rare; stomatal complexes cyclocytic, subsidiary cells very similar to non-modiied epidermal cells but more intensively staining, stomata oval to roundish, l × w = 24–37 (average 30) × 25–34 (average 29) µm, epidermal wall of guard cells often indistinct, front cavity 5–17 (average 11) µm long, almost reaching the poles, formed by thick and broad cuticular ledges, aperture short oval, polar I-thickenings ± well developed. R e m a r k s. Remarkable is the extremely irm texture causing a peculiar pattern of cracks that is unique among the plant remains of Rauenberg. So far, no putative taxonomic afinity can be suggested for this leaf. Dicotylophyllum oechsleri Kovar-Eder sp. nov. Pl. 7, ig. 17; Pl. 22, igs 1–4 Holotype 1952/454. designated h e r e. SMNS P S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. Dedicated to Harald and Annette Oechsler, who have collected the Rauenberg plant site systematically for many years. D i a g n o s i s. Coriaceous leaf with spiny marginal teeth; adaxial and abaxial cuticle leathery, anticlines undulate, stomatal complexes loosely spaced, anomocytic to indistinctly cyclocytic, subsidiary cells corresponding in shape and size to non-modiied cells, stomata elliptic to subcircular, front cavity short elliptic to roundish, prominent trichome bases of probably peltate glandular trichomes scattered. D e s c r i p t i o n. Leaf fragment lacking base and apex, texture coriaceous, l × w = 29 (? 55–60) × 18 (19–20) mm, laminar shape probably elliptic, margin coarsely serrate, teeth spiny, sinus rounded, basal and apical sides straight to concave; midvein slender, somewhat bent, secondaries craspedodromous, forking or sending abmedial veinlets into the tooth apices, further venation reticulate but exact details unclear. M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle thick with scattered simple trichome bases, margin collar-like thickened, surrounding cells short, radially arranged, the ? discshaped or globular head left a depression and a circular marginal ridge on the cuticle surface; adaxial cuticle: anticlines slender, deeply sinuous, cuticle surface distinctly, coarsely striate, non-modiied epidermal cells 29–54 (average 41) µm across; abaxial cuticle: anticlines thick, sinuous, non-modiied epidermal cells 37–54 (average 46) µm across, stomatal complexes anomocytic to indistinctly cyclocytic, subsidiary cells corresponding in shape and size to non-modiied cells, stomata elliptic to subcircular, l × w = 25–29 (average 26) × 18–26 (average 22) µm, guard cells less strongly cutinised than other cells, ledges well developed forming a broad elliptic to roundish front cavity, 9–13 (average 10) µm long. R e m a r k s. Gross morphologically this leaf is reminiscent of Ilex (Walther & Kvaček 2008) or Mahonia (Kvaček & Bůžek 1994). The cuticluar features do not conirm an afinity to Mahonia. The variability of the epidermal features of Ilex is wide (Baas 1975), making comparisons a challenge. Especially the stomata and type of stomatal complexes are diverse (Baas 1975, plate 2). Ilex aquifolia, for example, bears a thick adaxial cuticle with undulate anticlines and strong striation (Walther & Kvaček 2008) alike Dicotylophyllum oechsleri. In I. opaca the trichome bases resemble those encountered on our fossil (Walther & Kvaček 2008, pl. 6 ig. 9). Ilex foliage from Kreuzau (Germany, Miocene; Ferguson 1971) bears no resemblance gross morphologically, the adaxial cuticle is devoid of striae, and the stomatal complexes are clearly cyclocytic, but the stomata resemble our fossil in shape and size, as do the non-modiied epidermal cells of the abaxial cuticle. Trichome bases also occur but remained unigured and therefore cannot be compared. Ilex castellii Kvaček & Walther Unangemeldet Heruntergeladen am | 10.04.17 15:37 364 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 (Kvaček & Walther 1981) differs by, among other characters, the absence of striae adaxially and abaxially by having bigger stomata with a thickened outer margin and broad ledges forming the front cavity. I. knoblochii (Walther 1999) described from Kleinsaubernitz (Germany, Upper Oligocene) differs by its smaller undulation both in amplitude and wave length adaxially and abaxially, stomata arranged in groups, the presence of T-shaped polar thickenings, and the lack of trichome bases. Due to all these differences this leaf remains assigned to Dicotylophyllum only. Dicotylophyllum vesiculaeferens Kovar-Eder sp. nov. Pl. 7, igs 7, 8; Pl. 20, igs 1–5; Pl. 21, igs 1–3 Holotype 1952/310. designated h e r e. SMNS P Pa r a t y p e. SMNS P 1952/306. F u r t h e r m a t e r i a l. ? SMNS P 1952/414. S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. Referring to the vesicle-like, secretion-bearing structures (idioblasts). D i a g n o s i s. Narrow, entire-margined leaves, cuticles delicate with pentagonal to hexagonal pocket-like structures, single or, more often, occurring in groups or even adjacent to each other, each with a central plurilet mark, sometimes still bearing vesicle-like secretory bodies. D e s c r i p t i o n. Leaves coriaceous, laminar shape narrow, oblong to somewhat obovate, l × w = 52 (? 55) –64 (? 75) × 8–17 mm, l/w ratio = ? 4.4–6.8, entire-margined, margin thickened, base angle acute and narrow acute, base straight, apex angle narrow acute, apex straight or somewhat acuminate. M i c r o m o r p h o l o g y. Specimen SMNS P1952/310: Adaxial cuticle moderately thick, anticlines mainly slender, straight to inely undulate, partly pitted, non-modiied cells 17–25 (average 21) µm across; simple trichome bases scattered, trichome pore surrounded by somewhat radially elongated and slightly more strongly cutinised cells, single secretory structures (see abaxial cuticle) occur sporadically; abaxial cuticle delicate, anticlines of non-modiied epidermal cells almost straight to slightly bent, faintly visible if at all; cells polygonal, 14–24 (average 19) µm across, stomata weakly cutinised, dificult to discern, anomocytic, sometimes with tendency towards cyclocytic, roundish often asymmetric, stoma l × w = 11–18 (average 14) × 12–22 (average 17) µm, l/w ratio 0.7–0.9 (average 0.8), aperture short, slit-like or narrow oval, no thickened ledges developed, simple trichome bases as on adaxial cuticle, scattered; pentagonal to hexagonal pocket-like structures, 11–17 (14) µm in diameter, occurring in groups or even adjacent to each other, each with central tetralet to hexalet mark, sometimes still bearing vesicle-like secretory bodies; under SEM these structures appear elevated above the external cuticle surface, occasionally cuticle remnants of head still adherent. R e m a r k s. The most remarkable feature of these leaves is the mainly pentagonal to hexagonal pouches with a central asterix-like tetralet to hexalet mark, balloon-shaped secretory bodies, as well as cuticle remains of the head sometimes still adherent. The cuticle of specimen SMNS P 1952/306 is more poorly preserved and mainly hexagonal pouches are discernable. The material was studied by both light microscopy and SEM, but SEM studies were successful only on the external surface (prior to separating the cuticles). It remains unclear whether the secretory vesicles were secreted only below the cuticle or above it in a gland-like structure or both. These structures may have served to secrete salt or possibly other minerals. Complex salt glands do occur in different genera of the Chenopodiaceae, Tamaricaceae and Plumbaginaceae sensu Watson & Dallwitz (1992 onwards) (Salama et al. 1999). The plant assemblage from Rauenberg also yields articulate twig fragments with cuticles bearing pouches sometimes still yielding vesicle-like secretory bodies. The pouches are roundish or irregular in shape, however, not distinctly pentagonal to hexagonal. The twigs probably derive from the same plant species as the leaves because these structures are very extraordinary (see Cladites vesiculaeferens sp. nov.). Unangemeldet Heruntergeladen am | 10.04.17 15:37 365 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Dicotylophyllum ziegleri Kovar-Eder sp. nov. Pl. 7, ig. 6; Pl. 21, ig. 9; Pl. 22, igs 5–7 Holotype 1953/86. designated h e r e. SMNS P S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. In honour of the late Prof. Bernhard Ziegler, director of the Stuttgart State Museum of Natural History 1969–1994. D i a g n o s i s. Entire-margined, narrow obovate, shortly petiolate leaf; non-modiied cells large, abaxially large cyclocytic to somewhat actinocytic stomatal complexes, stomata narrow oval, ledges thickened, front cavity narrow spindle-shaped, almost reaching the poles, lacking polar T-pieces, aperture slit-like, discus- or bowl-shaped multicellular glands somewhat recessed in cuticle and overlapped by surrounding cells. D e s c r i p t i o n. Shortly petiolate, mediumthick leaf, laminar shape narrow obovate, l × w = 42 × 11 mm, l/w ratio = 3.8, petiole 2 mm long (? incomplete), almost 2 mm wide, basally widened; base cuneate, straight, base angle acute, apex angle acute, in the uppermost part obtuse, apex slightly retuse; midvein straight. M i c r o m o r p h o l o g y. Adaxial and abaxial cuticle well preserved, medium thick, smooth; anticlines slender but distinct, almost straight, non-modiied epidermal cells 34–61 (50) µm across adaxially and 26–45 (38) µm across abaxially; abaxial cuticle: stomatal complexes widely spaced, subparallel among each other, cyclocytic to slightly actinocytic, 5–8 subsidiary cells, similar in shape and size to non-modiied cells but less intensively staining, guard cells narrow oval, l × w = 36–50 (45) × 23–31 (27) µm, l/w ratio= 1.2–1.9 (1.7), epidermal wall of guard cell usually weakly cutinised, stomatal ledges strongly thickened, forming a narrow spindle-shaped front cavity, almost reaching the poles, aperture a faint slit; multicellular glands sunken and marginally overlapped by surrounding, marginally strongly thickened cells, diameter of pouches 34–47 µm (n = 5); SEM study yielded no further details. R e m a r k s. Due to its large non-modiied cells and stomatal complexes, along with the presence of epidermal secretory bodies, this leaf is reminiscent of Illicium, but cuticular studies of modern Illicium (Oh et al. 2003) indicate differences. Modern species were compared based on photographs provided by T. Denk, one of the coauthors of the mentioned paper. The cuticular features are rather consistent across modern species of Illicium. The stomatal complexes are brachyparacytic or more rarely (hemi)amphibrachyparacytic showing distinct polar T-pieces (Oh et al. 2003), while they are cyclocytic to actinocytic and lacking polar T-pieces in the heredescribed specimen. In Illicium the etherial oil cells are surrounded by 6–8 radially elongated cells, whereas in our specimen the surrounding cells are not elongated and the glands are sunken in pouches. In Illicium the anticlines are mainly undulate and the cuticle shows strong idiocuticular ornamentation, unlike Dicotylophyllum ziegleri in which the cuticle is smooth and the anticlines are straight. Illicium eocenicum (Jähnichen 1976), which is to be excluded from Illicium according to Oh et al. (2003), differs by the presence of polar T-pieces and the strong idiocuticular pattern from Dicotylophyllum ziegleri. Illicium geiseltalensis (Jähnichen 1976) – also to be excluded from Illicium (Oh et al. 2003) – differs by having smaller stomata with T-pieces, smaller non-modiied epidermal cells and secretory cells, and idiocuticular striation (Mai & Walther 1985). “Illicium” limburgense Kräusel & Weyland (Walther 1999) from Kleinsaubernitz is similar due to the presence of glands which appear sunken and overlapped by neighbouring cells and rather large stomata (though smaller than in D. ziegleri), but differs in leaf shape, petiole length, stoma shape and shape of the front cavity, and in having smaller glands and strong idiocuticular folding. Finally, there is no resemblance to the leaves of species XLIX in Ferguson (1971) that were previously described as Illicium liegelii (Weyland 1934). Dicotylophyllum sp. Pl. 7, ig. 18; Pl. 22, igs 8, 9 M a t e r i a l. SMNS P 1952/400. D e s c r i p t i o n. Petiolate, small leaf, rather complete, only apical quarter missing; petiole Unangemeldet Heruntergeladen am | 10.04.17 15:37 366 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 straight, 7 mm long (? complete), almost 1 mm wide, surface corrugated; lamina elliptic, texture medium thick, l × w = 25 (? 30) × 18 mm, l/w ratio about 1.7, base angle obtuse, base convex; margin entire, thickened, midvein straight, venation suprabasal acrodromous, basal secondaries arising almost oppositely about 1 mm above base, running at distance of 2–3 mm from the leaf margin, almost parallel to it and hardly tapering into the uppermost quarter of the lamina, no further veins visible. Magnoliaceae, have not been observed in this specimen. The systematic position of this leaf therefore remains ambiguous. M i c r o m o r p h o l o g y. Cuticles of both sides delicate, membranaceous, preserved in larger fragments, distinction between the two sides dificult due to faint stomata; anticlines indistinct, slender, almost straight, curved to slightly wavy, non-modiied cells 18–34 (average 22) µm across adaxially, 13–22 (average 19 µm) across abaxially, trichome bases present abaxially and ? adaxially, prominent, girdling cells ± distinct, roundish to pie-slicelike in shape, ± radially elongated, arranged concentrically around the trichome pore in one complete and sometimes a second incomplete circle, margin of trichome pore thickened, trichome pores 10–14 µm (n = 5) in diameter; abaxial cuticle: stomatal complexes only very rarely vaguely discernable, paracytic, broadly butterly-like due to broad lateral subsidiary cells, stomata ? sunken, 12–13 µm long (n = 2), stomatal ledges indistinct, outer front cavity narrow spindle-shaped, almost reaching the poles. P a r a t y p e s. SMNS P 1952/87, 371. R e m a r k s. Gross morphologically, Hamamelidaceae, e.g., Matudaea (Mai & Walther 1978), malvalean afinity, e.g., Byttneriopsis (Kvaček & Wilde 2010), or Lauraceae may be considered for this entire-margined, threeveined leaf. From the Daphnogene-type, this leaf differs by its rather delicate basal secondaries arising almost oppositely and very close to the base, running into the apical third of the lamina, hardly diminishing in width. Paracytic stomatal complexes occur in, for example, Ericaceae, Hamamelidaceae, or Magnoliaceae, while sunken ones may point to Lauraceae afinity. In the here-discussed specimen they are very small and only vaguely discernable if at all. Similar trichome bases do occur among Lauraceae, for example in shade leaves of Daphnogene (pl. 9, igs 4, 5). Oil-bearing idioblasts, as encountered in Lauraceae or Cladites D.H. Scott Cladites vesiculaeferens Kovar-Eder sp. nov. Pl. 7, igs 9–14, 19; Pl. 20, igs 6–9; Pl. 21, igs 4–8 H o l o t y p e. SMNS P 1952/416. M a t e r i a l. SMNS P 1952/104, 157; SMNS P 1953/105. NMA 2006-154/1497, NMA 20164/2179. SMNK 7778. S t r a t u m t y p i c u m. Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, early Oligocene. L o c u s t y p i c u s. Rauenberg, Baden-Württemberg, Germany. D e r i v a t i o n o m i n i s. Refering to the vesicle-like structures embedded in the cuticle. D i a g n o s i s. Telescope-like articulate axes, branched; cuticle: non-modiied epidermal cells short, ± in parallel rows, with ± clustered pouches, cuticle there thinner, sometimes with central plurilet mark, secretory bodies of very different size sometimes still adherent, balloon-shaped, originating as protuberances and embedded in pouches. D e s c r i p t i o n. Fragments of telescope-like articulate axes, branched (SMNS P 1952/87), segment length very variable, 2 mm (SMNS P 1952/87) to 26 mm (SMNS P 1952/371), segment width 2–6 mm. M i c r o m o r p h o l o g y. Cuticle very thick, anticlines strongly cutinised in older parts of branches but thin in apical/distal part; nonmodiied epidermal cells short, ± in parallel rows, pockets developed, cuticle there thinner, pockets ± clustered, variable in size, about 20–50 µm in diameter, sometimes with a central trilet to tetralet mark, secretory bodies of very different size, balloon-shaped, sometimes still adherent, originating as protuberances and embedded in pouches; in SEM view secretory bodies appear interconnected (pl. 21, ig 6); external surface of these structures in SEM view roundish to somewhat Unangemeldet Heruntergeladen am | 10.04.17 15:37 367 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 angular concavities with ± distinct centre and ± distinctly developed marginal cuticular rim (pl. 21, igs 7–8). and their vein-bearing wings are attached obliquely to the fruit body. Spiny branch R e m a r k s. These articulate stems, lacking stomata, supericially are reminiscent of rhizomes. The pocket-like structures of the cuticles are similar to those described for Dicotylophyllum vesiculaeferens. They differ in being more roundish or only weakly angular, and no complex structures elevated above the cuticle surface have been observed. The cuticle from a more distal part of the axis SMNS P 1952/416 implies that secretional structures developed from the cuticle and were embedded in it (pl. 20 ig. 9). Collection compartments separating the cuticle from the epidermis have been decribed from Tamaricaceae, Frankeniaceae, and Plumbaginaceae (Salama et al. 1999). The present structures possibly are similar. I suggest that the cross-like mark represents the region where the secretion was transferred to the stem surface. Depending on whether situated on the leaf or the stem, glands may be raised on the cuticle surface or sunken among epidermis cells, for example in Plumbaginaceae (Limoniastrum axillare, L. monopetalum) (Salama et al. 1999). The fragments of articulate branches and D. vesiculaeferens therefore probably derive from the same fossil species, and the complex secretory structures may represent salt glands (see also the section describing Dicotylophyllum vesiculaeferens). As for D. vesiculaeferens, the systematic afinity of the articulate stems remains open. A similar remain from Flörsheim has been described as a rhizome (Kvaček 2004a, rhizome B). This specimen was studied for comparison, but it is an impression only, devoid of organic material. Winged fruit or seed incertae sedis Pl. 6, ig. 3 M a t e r i a l. NMA 2006-113/1497. D e s c r i p t i o n. One-winged oval fruit or seed, l × w = 7 × 4 mm, wing membraneceous, l × w = 17 × 9 mm, dorsal line continuing rather straight to seed or fruit body, ventral line curved, no veins visible. R e m a r k s. This remain may be of coniferous origin. Fruits of Cedrelospermum are smaller, Pl. 7, ig. 15 M a t e r i a l. NMA 2006-23/1497. D e s c r i p t i o n. Fragment of axis, 23 mm long and about 2 mm wide, with spiny acute apex, two suboppositely originating branches, one of them complete, 52 mm long, terminating also in attenuate spine, the second branch incomplete. R e m a r k s. The main axis and the branches are cleary acute and attenuate spiny. This remain is similar to Fabaceae but spiny axes also occur in other families, for example in Rosaceae such as Crataegus or Prunus. RESULTS The plant assemblage of Rauenberg yielded 68 fossil species (Tab. 1). Three different types of marine algae were distinguished, among them representatives of the Phaeophyceae with aerocysts (Cystoseirites communis). Most remains, mainly leaves but also seed cones and other fructiications, derive from terrestrial plants. The taxonomic assignment of the terrestrial plant remains is based both on gross morphology and on cuticular features. Twenty-six families are documented, but nine taxa cannot be assigned to the family level. Fourteen fossil species of angiosperms are described for the irst time: Laurophyllum rauenbergense (Lauraceae), Myrica obliquifolia (Myricaceae), Distylium metzleri (Hamamelidaceae), ? Viscophyllum hendriksiae (? Loranthaceae), ? Berchemia altorhenana (? Rhamnaceae), ? Ternstroemites maritiae (? Pentaphyllaceae), Trachelospermum kelleri (Apocynaceae), Oleinites altorhenana, O. rauenbergensis (Oleaceae), Dicotylophyllum badense, D. oechsleri, D. vesiculaeferens, D. ziegleri, and Cladites vesiculaeferens. The diversity of Lauraceae (10) is highest, followed by Pinaceae (8), Arecaceae (4–5), Myricaceae (4), Cupressaceae (3, Taxodioideae 2, Cupressoideae 1), and Juglandaceae (3). All other families – Schizaeaceae, Zamiaceae, Doliostrobaceae, Platanaceae, ? Buxaceae, Fagaceae, Betulaceae, Betulaceae/Ulmaceae, ? Rhamnaceae, ? Fabaceae, Elaeocarpaceae, Salicaceae, Malvaceae, Hamamelidaceae, ? Loranthaceae, Unangemeldet Heruntergeladen am | 10.04.17 15:37 368 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 1. List of taxa and physiognomy. Abundance categories: I – 1 specimen, II – 2–5 specimens, III – 6–10 specimens, IV – 11–50 specimens, V – < 50 specimens. Laminar sizes in the Rauenberg lora: nanophyll 25–225 mm², microphyll 225– 2.025 mm², notophyll 2.025–4.500 mm², mesophyll 4.500–18.225 mm² (Webb 1955), calculated l × w × 2/3 (see Leaf Architecture Group 1999) Taxon Organ type Abundance Leaf margin Texture Laminar size Algae Cystoseyrites communis Thallites multiidus Laminarites latus I IV IV Ferns Lygodium kaulfussii Filicopsida sp. lealet lealet II I leaf I Cycads Ceratozamia loersheimensis Conifers leafy twig leafy twig leafy twig leafy twig fascicle fascicle fascicle fascicle fascicle fascicle cone cone I I II IV I II II I II I II I Actinodaphne pseudogermari Daphnogene cinnamomifolia leaf leaf I V entire entire coriaceous coriaceous Laurus abchasica leaf II entire II entire Doliostrobus taxiformis ? taxifomis Sequoia abietina Taxodium sp. Tetraclinis salicornioides Pinus (Strobus) sp. 1 Pinus (Strobus) sp. 2 Pinus (Strobus) sp. 3 Pinus (Strobus) sp. 4 Pinus sp. 5 Pinus sp. 6 Pinus (Pinus) cf. thomasiana Pinus engelhardtii Angiosperms Laurophyllum cf. kinkelinii var. glabrum leaf II entire Laurophyllum pseudoprinceps Laurophyllum rauenbergense leaf leaf IV IV entire entire moderately coriaceous moderately coriaceous moderately coriaceous coriaceous coriaceous Laurophyllum sp. A Laurophyllum sp. B Laurophyllum sp. C Palmacites lamanonis Sabalites major Phoenicites sp. ? Calamoideae gen. et sp. indet. Arecaceae gen. et sp. indet. Platanus neptuni leaf leaf leaf leaf leaf leaf leaf/petiole leaf leaf I I I IV II II I I V entire entire entire coriaceous coriaceous coriaceous coriaceous ? Buxus egeriana Eotrigonobalanus furcinervis f. haselbachensis Betula dryadum Betulaceae vel Ulmaceae gen. et sp. indet. leaf leaf I I basally entire, then serrate entire entire winged fruit leaf I I sharply ? double serrate Comptonia difformis leaf II membranaceous coriaceous Laurophyllum kinkelinii coriaceous coriaceous notophyll nanophyllnotophyll microphyll microphyll microphyll microphyll nanophyll, microphyll microphyll ? microphyll microphyll micro- to mesophyll nanophyll microphyll microphyll ? nanophyll, microphyll Unangemeldet Heruntergeladen am | 10.04.17 15:37 369 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 1. Continued Taxon Organ type Abundance Leaf margin Texture Laminar size moderately coriaceous moderately coriaceous delicate to moderately coriaceous membranaceous microphyll Myrica cf. lignitum leaf I Myrica longifolia leaf II Myrica obliquifolia leaf III Carya fragiliformis leaf I basally entire, then serrate almost entire to serrate basally entire, then minutely serrate sharply serrate Carya quadrangula Engelhardia orsbergensis fruit lealet II II serrate membranaceous nanophyll, microphyll Engelhardia macroptera ? Berchemia altorhenana involucrum leaf II I entire moderately coriaceous nanophyll ? Leguminocarpon sp. Sloanea artocarpites fruit leaf I I leaf II Populus germanica leaf II membranaceous membranaceous membranaceous notophyll Sloanea olmediaefolia wavy dentate, minutely toothed wavy dentate, minutely toothed wavy Craigia bronnii Distylium metzleri fruit leaf I I leaf calyx leaf leaf III I I II membranaceous coriaceous microphyll ? Viscophyllum hendriksiae Hydrangea microcalyx ? Ternstroemites maritiae Symplocos deichmuelleri apically remotely dentate entire Trachelospermum kelleri leaf II entire apically simple serrate entire coriaceous moderately coriaceous coriaceous Trachelospermum steiningeri leaf II entire Oleinites altorhenana leaf II entire Oleinites rauenbergensis Pungiphyllum cruciatum leaf leaf II I Dicotylophyllum badense Dicotylophyllum oechsleri leaf leaf I I Dicotylophyllum vesiculaeferens Dicotylophyllum ziegleri leaf leaf II I entire deeply lobate, lobes spiniform entire coarsely serrate, teeth spiny entire entire moderately coriaceous moderately coriaceous coriaceous ? membranaceous coriaceous coriaceous Dicotylophyllum sp. leaf I entire Cladites vesiculaeferens axis III Hydrangeaceae, ? Pentaphyllaceae, Symplocaceae, Apocynaceae, and Oleaceae – are represented by one or two species. The most abundant fossil species are Platanus neptuni, followed by Daphnogene cinnamomifolia, Tetraclinis salicornioides, Laurophyllum pseudoprinceps, and Palmacites lamanonis, while other species are represented by one or few remains only. Remains of ferns are extremely rare and not diverse: Lygodium kaulfussii, Filicopsida sp. Cycads are represented by a single lealet fragment of coriaceous moderately coriaceous moderately coriaceous microphyll microphyll notophyll microphyll, notophyll microphyll microphyll microphyll microphyll nanophyll, microphyll microphyll notophyll nanophyll microphyll ? microphyll microphyll microphyll microphyll microphyll Ceratozamia loersheimensis. Conifer diversity is rather high (12 species), of which Pinus (two ive-needled, one three-needled, three two-needled species, and two species based on cones) and Tetraclinis salicornioides (twig fragments) are well represented, whereas Taxodium sp. is documented by three, and Doliostrobus and Sequoia each by only a single foliated twig remain. Remarkably, in the Rauenberg assemblage, Fagaceae are almost absent except for a single leaf of Eotrigonobalanus furcinervis Unangemeldet Heruntergeladen am | 10.04.17 15:37 370 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 f. haselbachensis, and the record of Pentaphyllaceae is scarce and the remains ambiguous (? Ternstroemites maritiae). Fan palm remains are more common than those of feather palms, and ? Calamoideae are represented by a single strongly armed remain. Thirty-nine terrestrial woody angiosperm taxa are represented by simple entire-margined or mainly minutely serrate leaves. Two taxa, Pungiphyllum cruciatum and Dicotylophyllum oechsleri, show large spiny teeth. The leaf texture of more than two thirds of the dicots is coriaceous or moderately coriaceous, while less than one third have membranaceous leaves. The prevailing laminar size of the foliage is microphyll (67%), followed by notophyll (13%), nanophyll (8%), and mesophyll (1%). Cuticle preservation is reasonably good, but fungal infection is very common. The stomata are often masked by penetrating fungi, and the hyphae netwok disturbs the cuticular picture. DISCUSSION TAPHONOMY The lora of Rauenberg derives from mainly well-bedded to laminated clay to siltstones of marine origin (Bodenheim Formation). Terrestrial plant fossils are not condensed in layers or lenses but are found widely scattered, as is characteristic for marine sediments. Fragmentation of the plant material, resulting in leaves lacking apex or/and base, is common, indicating mechanical stress; relatively small-leaved taxa (predominant laminar size microphyll) comprise >70%. Tetraclinis salicornioides, one of the few abundant species, is represented mainly by single whorls, while articulate twigs are extremely rare. Although foliage of Platanus neptuni is by far the most common, not a single remain is reminiscent of a plane tree inlorescence or infructescence. These pecularities, along with high species diversity but low representation of most taxa, indicate transport from the growing site to the inal place of burial: that is, a highly allochthonous plant assemblage. Fungal infection of the leaf material may have started already prior to abcission; the often strong degree of infection points to a certain time span between leaf abcission and inal burial. The allochthonous deposition may explain the rarity of Fagaceae (a single leaf of Eotrigonobalanus furcinervis f. haselbachensis) and both the scarcity and ambiguity of the Pentaphyllaceae (? Ternstroemites maritiae). The extremely low representation of foliage of deciduous woody taxa may also partly relect mechanical stress during transport. Nevertheless, the absence of the respective winged fruits (except for a single, poorly preserved remain of Craigia bronnii, Betula dryadum and involucres of Engelhardia macroptera) may be interpreted as low representation of such taxa in the vegetation. The transport distance remains open, but the basinal Bodenheim Formation was deposited further away from the coast than its nearshore equivalent, the Alzey Formation (Grimm 2005). Rather shallow depositional depth was widely assumed by different authors (Weiler 1966, Micklich & Hildebrandt 2010, Micklich et al. 2009), but Grimm et al. (2002) argued for about 200 m water depth. Based on a comparison with modern relatives of the ish taxa, Maxwell et al. (2016) again propose moderately shallow conditions. The rich and diverse plant record is more supportive of the latter interpretation. ALGAE AND MONOCOTYLEDONS Band-like, thin, brownish, hardly carbonised fragments not showing venation represent marine algae (Laminarites latus, Thallites multiidus). Parallel-veined carbonised plant tissue was erroneously interpreted as “reeds” (Micklich & Hildebrandt 2010). Cuticular analysis revealed that such specimens represent highly fragmented palm foliage. A single remain turned out to represent the cycad Ceratozamia loersheimensis. No remains of reeds and sedges (Cyperaceae) or grasses (Poaceae) have been encountered. The presence of the sirenian “Halitherium” suggests the presence of seagrass meadows in the surrounding regions (Reich et al. 2015, Maxwell et al. 2016), but no seagrasses (Zosteraceae) have been detected among the rich fossil material. VEGETATION ANALYSIS AND RECONSTRUCTION Based on the autecology of modern relatives, autecological traits are inferred for the fossil species (Tab. 2). Probably more than 60% of the woody angiosperms were evergreen, while presumably deciduous taxa make up less Unangemeldet Heruntergeladen am | 10.04.17 15:37 371 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 than 20%; for about the same percentage it remains unclear whether they regularly shed their foliage. Deciduous representatives were Betulaceae vel Ulmaceae sp., Carya, possibly Engelhardia, Platanus neptuni, Populus germanica, and possibly Sloanea. The presence of winged fruits of Betula dryadum and Craigia bronnii as well as the calyx remain of Hydrangea microcalyx indicate higher diversity of deciduous taxa than indicated by leaf remains. Except for ferns, all terrestrial taxa reported (64) were woody. About 35% presumably were tall trees or trees. About the same percentage encompasses taxa that likely were small trees or shrubs, and >10% may represent lianas or climbers. Presumably anemophilous and zoophilous woody taxa are almost equally represented (about 40% each). Based on the hummingbirds recorded from Rauenberg (Mayr 2004), Apocynaceae, Loranthaceae, and Pentaphyllaceae may be considered to be possibly bird-pollinated taxa. Taxa with a presumably zoochorous fruit vector predominate over anemochorous ones (>40% versus about 35%) and about 10–15% were possibly dischorous or autochorous. The IPR vegetation analysis (Kovar-Eder & Kvaček 2008, Teodoridis et al. 2011b) enables a semiquantitative evaluation of the zonal vegetation type. This approach has been successfully tested and calibrated by application to modern vegetation in China and Japan (Teodoridis et al. 2011a). For the Rauenberg fossil lora, this analysis indicates broadleaved evergreen forests as zonal vegetation (BLE-component 49%, BLD-component 19%, SCL+LEG component 13–14%, conifers 12% of all zonal taxa) (Appendix 2). This result must be considered together with the allochthonous deposition of the Rauenberg assemblage (see section “Taphonomy”), which may have led to a certain overrepresentation of the BLE and SCL+LEG components at the cost of the BLD component. Nevertheless, the here-presented result is regarded as robust for two reasons: the threshold of the BLE component for broadleaved evergreen forests is >40% (Teodoridis et al. 2011a) and the number of presumably zonal taxa is high as compared to other fossil leaf assemblages. Sociological and physiognomical comparisons consistently indicate afinity to evergreen sclerophyllous broad-leaved forest (sensu Wang 1961; i.e. broad-leaved evergreen forest in the sense of IPR-vegetation analysis) as the most likely zonal vegetation type. Today, Fagaceae, Lauraceae, Pentaphyllaceae, and Magnoliaceae are dominant in this forest type and diversity is very high on the species level (Wang 1961). Numerous species (especially of Fagaceae) are closely allied, indicating rather recent diversiication. Such diversiication may be assumed due to the hybridisation idelity of modern oaks, which was discussed recently by Kovar-Eder et al. (2015). Evergreen taxa with coriaceous, ovate to lanceolate, either entire-margined or minutely serrate leaves are dominant in this forest type. Broad-leaved deciduous taxa and conifers do occur but are not important, except for some hardwoods in ecotones and also pines in successional series. Palms do occur but are less prominent than in rain forests (Wang 1961). Regarding composition, Rauenberg differs from evergreen sclerophyllous broad-leaved forest (sensu Wang 1961) in that only the diversity of Lauraceae is comparable, while Fagaceae and Pentaphyllaceae are hardly represented and Magnoliaceae are lacking. This may be linked to taphonomic bias caused by transport prior to deposition. This interpretation is supported by the presence of evergreen Fagaceae (3 species) and Pentaphyllaceae (2 species) in the nearby lora of Flörsheim (see section “The lora of Rauenberg in the European context”). In this context the evidence of Ceratozamia, Platanus neptuni, Sloanea, and Trachelospermum in Rauenberg is highly relevant. Ceratozamia is restricted to Central America (Mexico, Belize and Guatemala) today, where it occurs in mountains (800–1800 m) extending to lowlands; ecologically it ranges from tropical rain forests and less humid broad-leaved to seasonally dry forests with oaks and pines (Jones 1993). The relict species Platanus kerrii, which is regarded as an ecological equivalent of P. neptuni, occurs in Vietnam and Laos. Modern representatives of Sloanea, as opposed to the fossil species, thrive in SE China and Vietnam to Thailand (Kvaček & Hably 2001). Evidently, a comparison to modern vegetation types is limited in terms of composition. For Rauenberg, the attempt to reconstruct the vegetation is based on the autecology of modern relatives and a comparison with modern vegetation. The diversity and abundance of pines and palms indicate open near-coastal pine forests with palms on sandy soils. This Unangemeldet Heruntergeladen am | 10.04.17 15:37 372 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 2. Similar living relatives of the recorded terrestrial taxa, physiology derived from modern relatives: leaf physiology, growth form, primary pollination type, and primary dispersal mechanism; small tree < 15 m, tall tree > 15 m Similar living relative Taxon Inferred leaf physiology Growth form Primary pollination type Primary dispersal mechanism – – – – Ferns Lygodium kaulfussii Filicopsida sp. Lygodium ? – – climber – Cycads Ceratozamia loersheimensis Ceratozamia evergreen kuestleriana and allies small shrub zoophilous endozoochorous evergreen tall tree anemophilous anemochorous evergreen deciduous anemophilous anemophilous anemochorous anemochorous anemophilous anemochorous anemophilous anemophilous anemophilous anemophilous anemophilous anemochorous anemochorous anemochorous anemochorous anemochorous Conifers Doliostrobus taxiformis ? taxifomis Sequoia abietina Taxodium sp. Araucariaceae/ Taxodioideae Sequoia sempervirens Taxodium Tetraclinis salicornioides Tetraclinis articulata evergreen Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus Pinus evergreen evergreen evergreen evergreen evergreen tall tree tall tree small tree or shrub tall tree tall tree tall tree tall tree tall tree Actinodaphne ? Cinnamomum camphora Laurus nobilis and L. azorica evergreen tree or shrub entomophilous endozoochorous evergreen tall tree entomophilous endozoochorous evergreen small tree or shrub entomophilous endozoochorous Lauraceae evergreen tree entomophilous endozoochorous Ocotea Lauraceae Lauraceae Arecaceae Arecaceae Arecaceae evergreen evergreen evergreen evergreen evergreen evergreen tree tree tree tree tree tree shrub shrub shrub shrub entomophilous entomophilous entomophilous entomophilous entomophilous entomophilous endozoochorous endozoochorous endozoochorous endozoochorous endozoochorous endozoochorous Arecaceae evergreen tree or shrub entomophilous endozoochorous Arecaceae Platanus kerrii evergreen deciduous entomophilous anemophilous ? Buxus egeriana Buxus in SE-Asia evergreen tree or shrub tall tree shrub or small tree endozoochorous anemochorous auto-/endozoochorous Eotrigonobalanus furcinervis f. haselbachensis Betula dryadum Betulaceae vel Ulmaceae gen. et sp. indet. Comptonia difformis Trigonobalanus verticillatus Betula Betulaceae, Ulmaceae Comptonia peregrina evergreen tall tree anemophilous dyschorous deciduous tree (or shrub) anemophilous deciduous tree (or shrub) anemophilous deciduous anemophilous Myrica cf. lignitum Myrica ? evergreen anemophilous endozoochorous Myrica longifolia Myrica evergreen anemophilous endozoochorous Myrica obliquifolia Myrica evergreen anemophilous endozoochorous Carya fragiliformis Carya quadrangula Carya Carya shrub shrub or small tree shrub or small tree shrub or small tree tall tree tall tree anemochorous anemo-/dyschorous dyschorous anemophilous anemophilous dyschorous dyschorous anemophilous anemochorous (Strobus) sp. (5-needled) sp. (3 sp.) (2-needled) sp. (3-needled) (Pinus) cf. thomasiana engelhardtii Angiosperms Actinodaphne pseudogermari Daphnogene cinnamomifolia Laurus abchasica Laurophyllum kinkelinii – complex Laurophyllum pseudoprinceps Laurophyllum rauenbergense Laurophyllum sp. A–C (3) Palmacites lamanonis Sabalites major Phoenicites sp. ? Calamoideae gen. et sp. indet. Arecaceae gen. et sp. indet. Platanus neptuni Engelhardia orsbergensis/ macroptera deciduous deciduous deciduous, Engelhardia serrata, semiOreomunnea mexicana evergreen, evergreen or or or or tall tree Unangemeldet Heruntergeladen am | 10.04.17 15:37 373 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 2. Continued Similar living relative Taxon ? Berchemia altorhenana Berchemia ? Leguminocarpon sp. Fabaceae Sloanea artocarpites Sloanea olmediaefolia Populus germanica Sloanea in SE-Asia Sloanea sinensis Populus Craigia yunnanensis, C. kwangsiensis Craigia bronnii Inferred leaf physiology deciduous, evergreen deciduous, evergreen evergreen evergreen deciduous deciduous Growth form Primary pollination type climber or small to medium-sized entomophilous tree Primary dispersal mechanism endozoochorous tree entomophilous auto-/zoochorous tall tree tall tree tall tree entomophilous entomophilous anemophilous anemo- / entomophilous endozoochorous endozoochorous anemochorous ? autochorous small tree anemochorous Distylium metzleri Distylium evergreen ? Viscophyllum hendriksiae Loranthaceae evergreen Hydrangea microcalyx Hydrangea deciduous, evergreen shrub or small tree shrub, tree, liana shrub, small tree or climber evergreen tree or shrub tree or shrub liana liana tree, shrub, liana tree, shrub, liana ? ? ? ? ? ? endozoochorous anemochorous anemochorous endo-/anemochorous endo-/anemochorous ? ? ? entomo-/ ornithophilous entomophilous/ autophilous? entomo-/ornithophilous entomophilous entomophilous entomophilous zoochorous anemochorous Symplocos deichmuelleri Trachelospermum kelleri Trachelospermum steiningeri Pentaphyllaceae (former Theaceae) Symplocos Trachelospermum Trachelospermum Oleinites altorhenana Oleaceae Oleinites rauenbergensis Oleaceae Pungiphyllum cruciatum Dicotylophyllum badense Dicotylophyllum oechsleri Dicotylophyllum vesiculaeferens Dicotylophyllum ziegleri Dicotylophyllum sp. Cladites vesiculaeferens ? ? ? evergreen evergreen evergreen deciduous, evergreen deciduous, evergreen ? evergreen evergreen ? evergreen ? ? ? ? ? ? ? evergreen ? evergreen ? ? ? ? ? ? ? ? ? ? ? Ternstroemites maritiae interpretation is supported by the Alzey Formation, which represents the near-coastal sandy facies of the basinal Bodenheim Formation and its poorly preserved plant remains (Mallison 2002). Daphnogene cinnamomifolia and Myricaceae possibly were also represented in these forests, along with Ceratozamia loers heimensis, Carya and Engelhardia orsbergensis/macroptera, and Platanus neptuni. Coastal pine forests bound to sandy soils do occur in a wide variety and with numerous pine species along the eastern coast of the United States, although they are regarded mainly as secondary stands there (Okuda 1994). Cladites vesiculaeferens and Dicotylophyllum vesiculaeferens, which presumably derive from one plant species which was characterised by complex ? salt-excreting glands, thrived along the coast. Gallery forests along streams probably were characterised by associations of Platanus neptuni along with Daphnogene ? ? endozoochorous cinnamomifolia, Laurophyllum pseudoprinceps, Myricaceae, Populus germanica, Carya, Hydrangea microcalyx, and Craigia bronnii. Most other taxa represented by few or single remains may derive from zonal evergreen sclerophyllous broad-leaved forests. Some of the aforementioned probably were more generalists and may have thrived in these zonal forests as well, for example Daphno cinnamomifolia, Engelhardia orsbergensis/macroptera, L. pseudoprinceps, and Platanus neptuni. The ecological plasticity of Platanus neptuni was already discussed by Walther (1985). CLIMATE Climate estimates are based mainly on the modern occurrence of the zonal vegetation type, that is, southern regions of evergreen sclerophyllous broad-leaved forest (Wang 1961), which is assumed to be the closest analogon Unangemeldet Heruntergeladen am | 10.04.17 15:37 21–27 204–349 18–38 > 230 140.4 (104) 658.2 (239) 17.2–20.5 16–18 CA (Moraweck et al. submit.) CA Mainz Basin NP 23 (Pross et al. 1998) 25.0–25.9 25–28 12.6–13.6 6–10 1217–1322 1000–1300 1323.0 (497) 1300–1700 8–14 9.5 (2.6) 25.8 (1.7) 28–29 19–24 18.0 (1.3) Comparison with climate in southern regions of evergreen sclerophyllous broad-leaved forests in SE Asia and regions of most similar relevant modern relatives (this study) CLAMP (Moraweck et al. submit.) DMMP (mm) WMMP (mm) 3_dry (mm) 3_wet (mm) GSP (mm) MAP (mm) CMMT (C°) WMMT (C°) MAT (C°) Method applied for the lora of Rauenberg. Additionally, the climatic requirements of most similar living relatives of taxa such as Ceratozamia, Craigia, Platanus neptuni, and Sloanea are taken into consideration. Thus, the climatic conditions are most comparable to regions in Southeast Asia and America at around 20° N latitude today. In Southeast Asia, evergreen sclerophyllous broad-leaved forests extend mainly in regions with temperate, humid, summer hot climate (Cfa climate sensu Köppen 1936, Peel et al. 2007) or temperate, winter-dry, summer hot climate (Cwa climate) (Appendix 3). In the distribution area of Ceratozamia, in the southernmost parts of North America and northern Central America, tropical wet and tropical monsoon climate prevails. Floristic relations of the Rauenberg lora are, however, closer to Southeast Asia than to America. Therefore, the Rauenberg climatic conditions are interpreted as being comparable to the southern limits of Cfa climate, with possible tendencies towards a drier season in winter (Cw climate) and transition to tropical monsoon (Am) or tropical winter-dry (Af) climate. The climate may be further characterised as follows: Cfa climate in transition to Cwa (and Am or Af) climate, mean annual temperature (MAT) 19–24°C, mean annual precipitation (MAP) 1300–1700 mm, mean temperature of the warmest month (WMMT) 28–29°C, mean temperature of the coldest month (CMMT) 8–14°C, mean precipitation of the wettest month (WMMP) >230 mm, mean precipitation of the driest month (DMMP) 18–38 mm, wettest month between May and October, driest month between November and March. The warm period was wetter than the cold one. The upper limit of the derived MAT range appears somewhat high even when compared to existing ones for the Eocene (e.g., Kvaček et al. 2014, Zanazzi et al. 2007). For Rauenberg, CLAMP and the Coexistence Approach (CA) have also been applied (Moraweck et al. submit.) indicating somewhat lower MAT and WMMT values, while the CMMT estimates are fairly similar to that delivered by the modern occurrence of the zonal vegetation type (Tab. 3). CLAMP and CA signal a year-round growing season. MAP and growing season precipitation delivered by the different attempts overlap, but CA points towards lower MAP than that derived by the zonal vegetation type today. Most remarkably, especially the CA, the zonal vegetation type J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 3. Climate estimates derived by the application of different methods. MAT – mean annual temperature, CMMT – coldest month mean temperature, WMMT – warmest month mean temperature, MAP – mean annual precipitation, GSP – growing season precipitation, 3_wet – mean precipitation of the 3 wettest months, 3_dry – mean precipitation of the 3 driest months, WMMP – wettest month mean precipitation, DMMP – driest month mean precipitation 374 Unangemeldet Heruntergeladen am | 10.04.17 15:37 375 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 today and, to a minor degree, also CLAMP signal considerable seasonality in precipitation. For NP 23 in the Mainz Basin, Pross et al. (1998) estimated a Cfa-climate with lower values for temperature and precipitation (Tab. 3) but did not refer to possible seasonality. Those authors arrived at their results based on the CA applied mainly to the pollen record from the Bodenheim drillcore. These estimates were later accepted by Kvaček (2004a). The herepresented estimates for Rauenberg are based on a detailed taxonomic framework that enables correlation with a zonal vegetation type (southern regions of evergreen sclerophyllous broad-leaved forest), and the climate in the region of the occurrence of these forests today serves as a source for the climate proxy. The here-presented estimates for Rauenberg also differ from those presented by Grimm et al. (2011), who suggested a subtropicalmediterranean climate for the Alzey Formation, the coastal equivalent of the Bodenheim Formation. The basis for their assumption is unclear. Climate proxies for early Oligocene loras from North Bohemia and adjacent parts of Germany are also generally cooler than those presented in the current context for Frauenweiler (Kvaček et al. 2014). This is based on the high diversity of broad-leaved deciduous woody taxa there. These differences are not in conlict with the present results. Rather, they relect climatic differentiation within former Europe, where the climatic conditions along the western coast (and also southern coast) were warmer and probably exhibited a somewhat more pronounced seasonality of precipitation. The prevailing leaf size class microphyll may support this latter assumption of pronounced seasonality of precipitation. THE FLORA OF RAUENBERG IN THE EUROPEAN CONTEXT Diversiied early Oligocene loras are known from marine deposits in the Upper Rhine Graben (Bodenheim Formation) and from the Paratethys region (Tard Clay Formation) as well as from the early Miocene (Ebelsberg Formation (Rupp & Ćorić 2012; formerly “Älterer Schlier”, Upper Oligocene). As for Rauenberg, these plant assemblages are characterised by highly allochthonous deposition (Kovar 1982, Kvaček 2004a, Hably & Erdei 2015). Transport of the plant debris from the growing site to the burial place caused sorting and selection on the one hand, and on the other hand may account for a mixture of plant remains from presumably very different habitats. In contrast, early Oligocene taphocoenoses from northern Bohemia and adjacent parts of Germany (mainly Saxony) derive mainly from lacustric deposits in the adjacent volcanic hinterland (volcanic loras sensu Kvaček & Walther 2001) or were deposited in coastal lowlands (Leipzig Embayment, Mai & Walther 1978). Sorting and selection due to transport was less severe, e.g., Seifhennersdorf (Walther & Kvaček 2007), Bechlejovice (Kvaček & Walther 2004), Haselbach Floristic Complex (Kunzmann & Walther (2012). These general characteristics are noteworthy for the following comparisons. Floristic complexes (sensu Mai 1995) aim to characterise the development of the forest lora across Europe. The lora of Rauenberg, however, is compared here mainly to individual loras instead of loristic complexes. This provides a better basis to further develop the scheme of loristic complexes, especially regarding age correlation. The following comparisons are performed regionally, followed by stratigraphic position. FLORAS FROM THE RHINE GRABEN AND THE PARATETHYS REGION Flörsheim near Mainz, Germany, Upper Rhine Graben Unsurprisingly, the loras from Rauenberg and Flörsheim, both very diverse, deriving from the Bodenheim Formation and dated to nannoplankton zone NP 23, show closest similarities compared to the remaining European record. Platanus neptuni and Daphnogene cinnamomifolia are the most common species at both sites. Rauenberg and Flörsheim share 26 fossil species of 16 families (Tab. 4). The cycad Ceratozamia loersheimensis was irst described from Flörsheim. Remarkable is, among others, the presence of the following taxa in Rauenberg (absent in Flörsheim): Doliostobus taxiformis, Engelhardia orsbergensis and E. macroptera, Myrica obliquifolia (rather common in Rauenberg), Distylium, and Sloanea olmediaefolia. Fagaceae and Pentaphyllaceae (three species each, some of them abundant) play a much more important role in Flörsheim than in Rauenberg, and the record Unangemeldet Heruntergeladen am | 10.04.17 15:37 376 from Flörsheim includes the following taxa not discovered in Rauenberg: Chamaecyparites hardtii (Göppert) Endlicher, Laurophyllum acutimontanum Mai, Daphnogene engelhardtii Kvaček, 2 species of Andromediphyllum, Symplocos oligocaenica Kvaček, Oleinites pachyphyllus Kvaček, and Smilax weberi Wessel in Wessel & Weber. As in Rauenberg, also in Flörsheim broad-leaved laurophyllous species with entire or minutely serrate margin prevail over presumably deciduous modern ArctoTertiary taxa (sensu Kvaček 1994). Among the latter, Betulaceae are represented by leaves of three genera (Alnus, Carpinus, Ostrya) in Flörsheim, each with one species, but Betula, which is documented by a winged fruit from Rauenberg, is not among them. Craigia and Hydrangea have been detected only in Rauenberg. Mai (1995) established the Floristic Complex Nerchau-Flörsheim (see section “The lora of Rauenberg in the context of the concept of loristic complexes”). Orsberg, Siebengebirge, Rheinland-Plaz (Germany), southern margin of the Lower Rhenish Embayment Based on the dating of the nearby site Rott (mammal Paleogene zone MP 30), this recently revised leaf and fruit lora is thought to be late Oligocene/early Miocene in age (Winterscheid & Kvaček 2014). The taphocoenosis from the lignites (mainly fructiications) yields an impoverished Younger Mastixioid Flora (sensu Mai 1995), while the younger taphocoenosis from bituminous oil shales represents the remnants of riparian and swamp forests. Among conifers, Orsberg and Rauenberg share Sequoia abietina and Tetraclinis salicornioides. The swamp element Glyptostrobus europaeus (Brongniart) Unger, represented by twigs and cones in Orsberg, underpins the differences in facies to Rauenberg. In addition to taxa shared with Rauenberg (Tab. 4), leaves of Dombeyopsis lobata Unger, present in Orsberg, and a single fruit of Craigia bronnii from Rauenberg represent a single Malvaceae species. Laurophyllum acutimontanum and L. haemisphaericum (Roselt & Schneider) Winterscheid & Kvaček occur only in Orsberg. L. sp. 2 from Orsberg resembles the leaf SMNS P 1953/76 included in Daphnogene cinnamomifolia from Rauenberg. Leaf taxa not present in Rauenberg include: Acer, Alnus, Corylopsiphyllum celtifolium (C.O. Weber in P. Wessel & C.O. J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Weber) Winterscheid & Kvaček, Vaccinioides cf. lusatica (Litke) Kvaček & Walther, Pentaphyllaceae (3 species), Vitaceae (2), and others of cryptic afinity. Tard Clay Formation, Hungary, Paratethys region The marine Tard Clay Formation is dated to the nannoplankton zones NP 21–23, i.e. early Oligocene. Based on maceral, biomarker as well as stable isotope composition, facies changes from marine to brackish and reverse were traced (Bechtel et al. 2012). Based on the preservation type, most terrestrial plant taxa are known by gross morphology only, and no complete inventory of revised taxa is available. This hampers a full comparison (Hably & Erdei 2015). Apart from the presence and abundance of Platanus neptuni, Daphnogene cinnamomifolia and Tetraclinis salicornioides, Rauenberg and the Tard Clay Formation share the occurrence of Ceratozamia loersheimensis, Doliostrobus taxiformis, and Sloanea olmediaefolia (Tab. 4). None of the Lauraceae (Laurophyllum) identiied based on cuticles from the Tard Clay Formation occur at Rauenberg. Contrary to Rauenberg, Eotrigonobalanus furcinervis is very common. Taxa characteristic of the Tard Clay Formation such as Ziziphus zizyphoides (Unger) Weyland, Cedrelospermum aquense (Saporta) Saporta, Raskya vetusta (Ettingshausen) Manchester & Hably, Kydia kraeuselii (Rásky) Hably and the diversity of putative legumes distinguish this early Oligocene lora from the Hungarian part of the Paratethys region from that of the Upper Rhine Graben (Rauenberg and Flörsheim). Linz, Austria, Central Paratethys This lora derives from the Ebelsberg Formation, formerly “Älterer Schlier” (Kovar 1982). Nannoplankton indicates zone NN 1, i.e. Lower Miocene (Rupp & Ćorić 2012). Similarly to Rauenberg, the terrestrial plant remains occurred widely scattered in the sediment and were associated with both marine and terrestrial organism groups (Kovar 1982). In both regions, near-coastal sandy sediments are developed (Alzey Formation in the Upper Rhine valley, Linzer Sande along the Paratethys coast around Linz). Rauenberg and Linz share, among others, the diversity of pines and fan palms (Tab. 4), indicating near-coastal Unangemeldet Heruntergeladen am | 10.04.17 15:37 377 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 pine forests on sandy soils, and the abundance of Platanus neptuni, Daphnogene cinnamomifolia and Tetraclinis salicornioides. Except for numerous taxa present solely in Rauenberg, major distinguishing features are the presence of Cunninghamia miocenica Ettingshausen, Trigonobalanopsis rhamnoides (Rossmaessler) Kvaček & Walther, Acer, Fagus, Betulaceae (2), and Ulmaceae (2) only in Linz. Some of these discrepancies presumably relect differences in age, such as the absence of Doliostrobus in Linz or the absence of Cunninghamia in Rauenberg. FLORAS FROM THE LEIPZIG EMBAYMENT (GERMANY) AND THE VOLCANIC FLORAS FROM THE ČESKÉ STŘEDOHOŘÍ MOUNTAINS, NORTH BOHEMIA (CZECH REPUBLIC) AND SAXONY (GERMANY) The Floristic Complex Haselbach, Leipzig Embayment, Weisselster Basin, Saxony, Sachsen-Anhalt, Germany The Floristic Complex Haselbach (Mai & Walther 1978) comprises a suite of mainly parautochthonous assemblages relecting different, prevailingly intrazonal habitats such as swamp forests, riparian forests, pioneer vegetation and luvio-lacustric aquatic assemblages of the extended near-coastal lowland in the Leipzig Embayment (Kunzmann & Walther 2012). The stratigraphic assignment to the Lower Oligocene is based on palynological data (occurrence of Boehlenpollis hohlii Krutzsch; Krutzsch 2011). Plants of lacustric, swampy, and luvial settings are highly diversiied both in the leaf and fruit record, whereas zonal taxa are mainly restricted to the fruit record (Mai & Walther 1978). Conifers are very diverse, including numerous taxa of wetland and swamp environments as well as Tsuga and Cephalotaxus, while Doliostrobus, which is present in Rauenberg, has not been recorded (Tab. 4). Generally, broad-leaved deciduous woody angiosperms are diversiied in the Floristic Complex Haselbach, while neither Alnus, Carpinus, Salix, Acer or Liquidambar, to mention some, have been encountered in Rauenberg. Shared woody angiosperms between Rauenberg and the Floristic Complex Haselbach are mainly restricted to coriaceous ones that probably preferred intrazonal settings. This includes fan palms, Daphnogene cinnamomifolia, Laurophyllum pseudoprinceps, Platanus neptuni, and Populus germanica. The latter two do occur in both regions abundantly, but Eotrigonobalanus furcinervis is common only in the Haselbach Floristic Complex. Both loras further share the presence of Distylium, Symplocos and palms of putative calamoid afinity, although all are represented by different species. Lauraceae are much less diversiied in the Floristic Complex Haselbach (4 species) than in Rauenberg (10). Remarkable is the absence of Ceratozamia, Craigia, Hydrangea, Sloanea, Trachelospermum, and Oleaceae from the Floristic Complex Haselbach. Bechlejovice, České středohoří Mountains, Czech Republic This fossil leaf assemblage derives from diatomites deposited in a lake that had developed in a volcanogenic environment. Radiometric dating of a tephrite intrusion (26.8 ±1.34 Ma) indicates an older age of this lora than this date. Due to the presence of archaic taxa Kvaček & Walther (2004) consider this lora to derive from the basal part of the Lower Oligocene and assign it to the Floristic Complex Haselbach – Valeč. The plant remains document lora and associations in the nearby lake surroundings. The lack of cuticles hampers loristic comparisons. In Bechlejovice, the only available conifer is Torreya. Among angiosperms, Platanus neptuni does occur but it is extremely rare. Bechlejovice and Rauenberg share Carya, Craigia bronnii, Comptonia difformis, Laurophyllum pseudoprinceps, Pungiphyllum cruciatum, Sloanea artocarpites, and possibly also Zamiaceae (Tab. 4). Otherwise, Bechlejovice differs by the high diversity of broad-leaved deciduous taxa (4 genera and at least 5 species of Betulaceae, at least 4 maple species, etc.) compared to broad-leaved laurophyllous ones. It also differs by the presence of conservative taxa such as Platanus schimperi (Heer) Saporta & Marion, Sterculia crassinervia (Ettingshausen) Procházka & Bůžek, and Haemanthophyllum. Kundratice, České středohoří Mountains, Czech Republic The lora of Kundratice was deposited in a volcanic lake. Radiometrical dating of the base of the superimposed basalt provided an Unangemeldet Heruntergeladen am | 10.04.17 15:37 378 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 4. Floristic comparisons. Taxa in the lora of Rauenberg compared to Oligocene European sites Taxa in Rauenberg Microthyriaceae Cystoseirites communis Thallites multiidus Laminarites latus Lygodium kaulfussii Filicopsida sp. Ceratozamia loersheimensis Doliostobus taxiformis Sequoia abietina Taxodium sp. Tetraclinis salicornioides Pinus (Strobus) sp. 1 Pinus (Strobus) sp. 2 Pinus sp. 3 Pinus sp. 4 Pinus sp. 5 Pinus sp. 6 Pinus (Pinus) cf. thomasiana Pinus engelhardtii Actinodaphne pseudogermari Daphnogene cinnamomifolia Laurus abchasica Laurophyllum kinkelinii Laurophyllum cf. kinkelinii glabrum Flörsheim (Kvaček 2004a) Orsberg (Winterscheid & Kvaček 2014) Tard Clay (Hably 1979, Hably & Erdei 2015) + C. altoaustriacus + + + + + + + + + + + + S. langsdorii Taxodium dubium + and T. brachyodon ? Calamoideae gen. et sp. indet. Arecaceae gen. et sp.indet. Platanus neptuni ? Buxus egeriana Eotrigonobalanus furcinervis f. haselbachensis Betula dryadum Betulaceae vel Ulmaceae gen. et sp. indet. Comptonia difformis Myrica cf. lignitum Myrica longifolia Myrica obliquifolia Carya fragiliformis + + + P. palaeostrobus + T. dubium + P. palaeostrobus + Pinus sp. (3) P. div. sp. + + + + + + + + D. div. sp. D. sp. + + + + Laurophyllum + pseudoprinceps Laurophyllum rauenbergense as L. cf. villense Laurophyllum sp. A Laurophyllum sp. B Laurophyllum sp. C Palmacites lamanonis + Sabalites major Phoenicites sp. Linz, Austria (Kovar 1982) Haselbach Complex (Mai & Walther 1978, Kunzmann & Walther 2012) Sabal major Sabal raphifolia as ? Phoenicites sp. Calamus daemonorops + + E. furcinervis + as C. schrankii M. lignitum + + B. egeriana + M. lignitum C. acutiloba + E. furcinervis C. acutiloba M. lignitum cf. C. acutiloba + Carya sp. Unangemeldet Heruntergeladen am | 10.04.17 15:37 379 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 4. Continued Bechlejovice (Kvaček & Walther 2004) Kundratice (Kvaček & Walther 1998) Seifhennersdorf (Walther & Kvaček 2007) Hammerunterwiesenthal (Walther 1998) Markvartice (Bůžek et al. 1976, Kvaček & Walther 2001) SuleticeBerand (Kvaček Kleinsaubernitz & Walther (Walther 1999) 1995) ? Zamiaceae + T. dubium + + + + + + + + + P. palaeostrobus P. sp. + L. cf. pseudoprinceps L. cf. pseudoprinceps + + + L. cf. pseudoprinceps as D. lanceolata + 0 + + + + + + Sabal cf. lamanonis Sabal sp. Arecaceae + + + cf. E. furcinervis B. sp. fruits, B. buzekii + + B. kleinsaubernitzensis + C. cf. difformis C. cf. serrifolia, C sp. + M. lignitum C. serrifolia + C. cf. serrifolia C. serrifolia Unangemeldet Heruntergeladen am | 10.04.17 15:37 380 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 4. Continued Taxa in Rauenberg Carya quadrangula Flörsheim (Kvaček 2004a) Linz, Austria (Kovar 1982) + Sloanea olmediaefolia Populus germanica Craigia bronnii + Haselbach Complex (Mai & Walther 1978, Kunzmann & Walther 2012) + + + Sloanea artocarpites Trachelospermum kelleri Trachelospermum steiningeri Oleinites altorhenana Oleinites rauenbergensis Pungiphyllum cruciatum Dicotylophyllum badense Dicotylophyllum oechsleri Dicotylophyllum vesiculaeferens Dicotylophyllum ziegleri Dicotylophyllum sp. Cladites vesiculaeferens Winged fruit or seed incertae sedis Spiny branch Tard Clay (Hably 1979, Hably & Erdei 2015) + Engelhardia orsbergensis Engelhardia macroptera ? Berchemia altorhenana ? Leguminocarpon sp. Distylium metzleri ? Viscophyllum hendriksiae Hydrangea microcalyx ? Ternstroemites maritiae Symplocos deichmuelleri Orsberg (Winterscheid & Kvaček 2014) + + + + + + + Dombeyopsis lobata + + + + + + age of 32.7 ± 0.8 Ma (Bellon et al. 1998). Thus, the lora is presumably closer in age to that of Rauenberg than that of Seifhennersdorf (see below). Conifers are relatively diverse, represented by Cephalotaxus, Taxus, Tetraclinis, and Torreya. Tetraclinis is the only shared one between Rauenberg and Kundratice (Tab. 4). Neither Pinus nor Taxodioideae are recorded from Kundratice. Deciduous angiosperms comprise Acer (at least 5 species), Alnus, Betula, Carpinus, Craigia, Cercidiphyllum, Ostrya, Tilia, Ulmus, and Zelkova, of which Kundratice and Rauenberg share only Betula, Carya, and Craigia. Of ive species of Lauraceae in Kundratice and 10 in Rauenberg, both sites have only Daphnogene cinnamomifolia and possibly Laurophyllum pseudoprinceps in common. They further share Pungiphyllum cruciatum, Sloanea artocarpites, and Symplocos deichmuelleri. Both sites are characterised by the absence (Kundratice) or almost absence (Rauenberg) of Fagaceae. The Floristic Complex Seifhennersdorf-Kundratice (Kvaček & Walther 1998) is characterised as Mixed Mesophytic Forest vegetation with considerable diversity of broad-leaved deciduous taxa, i.e. modern Arcto-Tertiary elements. Unangemeldet Heruntergeladen am | 10.04.17 15:37 381 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Table 4. Continued Bechlejovice (Kvaček & Walther 2004) Kundratice (Kvaček & Walther 1998) C. costata + L. div. sp. cf. S. artocarpites as Icaciniphyllum artocarpites + and Dombeyopsis lobata + and Dombeyopsis lobata Seifhennersdorf (Walther & Kvaček 2007) Hammerunterwiesenthal (Walther 1998) C. cf. quadrangula + + + SuleticeBerand (Kvaček Kleinsaubernitz & Walther (Walther 1999) 1995) + + + Leguminosae gen. et sp. indet. as + as Dicotylophyllum Icaciniphyllum sparsidentatum artocarpites L. div. sp. + and Dombeyopsis lobata Markvartice (Bůžek et al. 1976, Kvaček & Walther 2001) + + as Pteleaecarpum europaeum + + + as Dicotylophyl- as Dicotylophyllum lum deichmuelleri deichmuelleri + + Seifhennersdorf, Upper Lusatia, Saxony, Germany The lora of Seifhennersdorf (Kvaček & Walther 2007) derives from diatomites that developed in a maar lake. Radiometric dating of the overlying basalts provided an age of 30.7 ± 0.7 Ma (Bellon et al. 1998). Thus, this lora is somewhat younger than that of Kundratice and Rauenberg. Taxa of aquatic and swamp associations distinguish the lora from Seifhennersdorf from that of Rauenberg. The high diversity of deciduous woody angiosperms, such as Acer (at least 7 species), Betulaceae (4 genera), and Rosaceae (4 species) distinguishes this lora strongly from + that of Rauenberg. Both sites, however, share Carya, Craigia bronnii and Hydrangea microcalyx (Tab. 4). Seifhennersdorf is characterised by the scarcity of Eotrigonobalanus furcinervis, as is Rauenberg, and the occurrence of Quercus lonchitis Unger. Lauraceae are less diverse in Seifhennersdorf and both sites share only Daphnogene cinnamomifolia and Laurophyllum pseudoprinceps. Remarkable is the presence of putative Oleaceae, although they are represented by different species in Seifhennersdorf (Oleinites hallbaueri (Mai) Sachse, O. maii (Bůžek, Holý & Kvaček) Sachse) and Rauenberg (O. altorhenana, O. rauenbergensis). Platanus neptuni also occurs in Seifhennersdorf but is less abundant Unangemeldet Heruntergeladen am | 10.04.17 15:37 382 than in Rauenberg. Besides Kundratice, Seifhennersodrf is the type locality of the Floristic Complex Seifhennersdorf-Kundratice (Kvaček & Walther 1998). Hammerunterwiesenthal, Saxony, Germany Age constraints of this lora were provided by radiometric dating of the basaltic tufites there: 30.48 ± 0.41 Ma (Walther 1998). Deposited in a maar lake, this plant assemblage is less diverse (25 taxa). With its extremely low diversity of conifers – Pinus, Tetraclinis salicornioides, both present in Rauenberg (Tab. 4) – and diversity of broad-leaved deciduous taxa – Acer (4 species), Alnus, cf. Carpinus, cf. Cercidiphyllum, Craigia, Hydrangea, Vitis – this lora resembles most of the other volcanic loras of this region but differs from the assemblage of Rauenberg. Hammerunterwiesenthal and Rauenberg share only Craigia and Hydrangea. Out of three laurels, Daphnogene cinnamomifolia and probaly Laurophyllum pseudoprinceps occur at both sites, while L. acutimontanum is restricted to Hammerunterwiesenthal. Trigonobalanopsis rhamnoides represents the only Fagaceae at this site. Fan palm indings are generally rare in the volcanic loras; a single one derives from Hammerunterwiesenthal. Markvartice, České středohoří Mountains, Czech Republic The stratigraphic position of this lora, which derives from tuffaceous diatomite and ash tufite, is somewhat ambiguous. Based on regional geology, loristic composition and relation to other loras, such as that from Seifhennersdorf, this lora is estimated to be somewhat younger (Bůžek et al. 1976). Kvaček & Walther (2003) assign this site to the Děčín Formation which is regarded mainly late Oligocene in age. Taxodioideae are hardly documented, pines are absent, Taxus and Torreya are represented by single specimens. The only common conifer is Tetraclinis salicornioides. The assemblage from Markvartice differs from others of this region by the prevalence of evergreen laurophyllous taxa. The diversity of Lauraceae is almost as high as in Rauenberg. Both sites, however, share only Daphnogene cinnamomifolia and Laurophyllum pseudoprinceps (Tab. 4). Similarly to Rauenberg, D. cinnamomifolia, L. pseudoprinceps, and Platanus neptuni are very abundant and J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 T. salicornioides is among the common species. Among deciduous taxa, Rauenberg and Markvartice share Betula dryadum, Carya, and Craigia bronnii. Markvartice, however, includes further broad-leaved deciduous taxa such as Acer (at least 3 species), Cercidiphyllum, Rosa, and Ulmus not documented in Rauenberg. Both sites share the presence of Sloanea artocarpites and putative Oleaceae, although different species (Oleinites maii in Markvartice, O. altorhenana and O. rauenbergensis in Rauenberg). Documented in Rauenberg (solely by a single leaf of Eotrigonobalanus f. furcinervis haselbachensis), Fagaceae records are absent at Markvartice. Mai (1995) and Kvaček & Walther (2001) assign this assemblage to the Floristic Complex Nerchau-Flörsheim. Suletice-Berand, České středohoří Mountains, Czech Republic Radiometric dating delimits the age of this lora at between 28.3 ± 1.4 Ma and 19.8 ± 0.5 Ma (Bellon et al. 1998). This taphocoenosis was embedded in diatomites that developed in a volcanic lake setting. The comparison is biased because cuticles are rarely preserved. Conifers are not diversiied; neither pines nor Taxodioideae occur. Only Tetraclinis salicornioides is rather frequent (Tab. 4). Among Lauraceae (at least 4 species), Laurophyllum acutimontanum, which is not recorded from Rauenberg, is the most common one, followed by Daphnogene cinnamomifolia and Laurophyllum pseudoprinceps. Platanus neptuni is abundant. Broad-leaved deciduous taxa are less diversiied than for example in Seifhennersdorf or Kundratice. Among these are Acer (at least 3 species), Carpinus, Celtis, Ostrya, Rosa, and Zelkova, none of which have been detected in Rauenberg. Suletice-Berand and Rauenberg, however, share Carya, Craigia bronnii, Engelhardia orsebrgensis/macroptera, Hydrangea microcalyx, and Sloanea artocarpites. Putative Oleaceae are represented by different Oleinites species at both sites. Kvaček & Walther (1995) assign this assemblage to the Floristic Complex Nerchau-Flörsheim. Kleinsaubernitz, Saxony (Germany) This rich lora derives from drill cores through maar lake deposits in which the plantbearing sediments are 270 m thick. Based on regional geology, Kleinsaubernitz was Unangemeldet Heruntergeladen am | 10.04.17 15:37 383 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 correlated to the Upper Oligocene (Walther 1999). This was rendered more precisely to Spore-Pollen-Paleogene zone 20G (Goth et al. 2003), corresponding to the lower Upper Oligocene (Krutzsch 2011). Conifers are very diverse (Pinaceae 3 species, Cupressaceae 5, Taxaceae 1), but Kleinsaubernitz and Rauenberg share only ive-needled Pinus, Sequoia abietina, Taxodium, and Tetraclinis salicornioides (Tab. 4). Angiosperms include laurophyllous taxa as well as “modern” Arcto-Tertiary ones, each group represented about 50%. Contrary to Rauenberg, Fagaceae are very diverse (8 species), among them the archaic Eotrigonobalanus furcinervis f. haselbachensis as well as Trigonobalanopsis rhamnoides and Fagus saxonica Kvaček & Walther. The presence of further archaic taxa such as Majanthemophyllum petiolatum Weber, together with taxa known from Miocene deposits such as “Illicium” limburgense Kräusel & Weyland and Illipophyllum thomsonii Kräusel & Weyland, are unique. Deciduous taxa are rather diverse (e.g. Acer, Alnus, Betula, Carpinus, Celtis, Craigia, Ulmus) but Kleinsaubernitz and Rauenberg share only the presence of Betula and Craigia bronnii. Further taxa occurring at both sites are Comptonia difformis, Sloanea artocarpites, and Distylium, the latter being represented by different species. Among the loristic record from the volcanic region of North Bohemia and adjacent parts of Saxony, Kleinsaubernitz is quite peculiar, which encouraged Walther (1999) to establish the Floristic Complex Kleinsaubernitz. SUMMARISING THE COMPARISON OF RAUENBERG WITH THE FLORAS FROM THE LEIPZIG EMBAYMENT AND THE VOLCANIC FLORAS FROM SAXONY AND THE ČESKÉ STŘEDOHOŘÍ MOUNTAINS From a purely loristical viewpoint the following characteristics are noteworthy: Daphnogene cinnamomifolia, Laurophyllum pseudoprinceps, Platanus neptuni, and Tetraclinis salicornioides are the taxa which are present and mostly also very abundant at almost all sites. Additionally, Rauenberg and all volcanic loras share the presence of Craigia bronnii, which is usually an accessory element. Rauenberg is distinct by the presence of Ceratozamia loersheimensis, Doliostrobus taxiformis, Myrica longifolia, and Sloanea olmediaefolia, which indicate an afinity to loras from the Paratethys realm. Otherwise, M. longifolia occurs only in the Floristic Complex Haselbach. Generally, Lauraceae (10 species), Myricaceae (4 ) and Arecaceae (4–5) are more diverse in Rauenberg than in any of the compared loras. The diversity and relative abundance of pines at Rauenberg, versus their absence or scarcity in the volcanic loras, indirectly supports the interpretation of near-coast pine forests in Rauenberg. At all compared sites from this part of Central Europe, Acer is present and more or less species-diverse, while there is not even a single record from Rauenberg. The high diversity of further “modern” ArctoTertiary elements distinguishes all these sites from Rauenberg. The diversity of taxa with simple, coriaceous, entire-margined or minutely toothed, probably evergreen leaves in Rauenberg is not only higher than in the loras from the region of Saxony, Saxony-Anhalt and North Bohemia, but many of them have never been recorded there. The latter two arguments (differences in diversity of “modern” Arcto-Tertiary elements and in prevailing leaf physiognomy) as well as the presence of different palms in Rauenberg explain the different estimated zonal forest types, namely mixed mesophytic forest or transition from mixed mesophytic to broad-leaved evergreen forest for most of the loras from North Bohemia, Saxony and Saxony-Anhalt, but broad-leaved evergreen sclerophyllous forest for Rauenberg. The loristic complex Haselbach repeatedly was interpreted as mixed mesophytic forest (Mai & Walther 1978, 1985, Mai 1995). Only recently broadleaved evergreen forest was predicted based on the IPR vegetation analysis (Kvaček et al. 2014). The percentage of the BLE component of zonal woody taxa is, however, lower there (42.5%) than in Rauenberg (48.6%). Regarding age correlations, the plant assemblages from Kundratice and Seifhennersdorf are probably closest to Rauenberg, but their loristic match with Rauenberg is lower than that of Markvartice, whose age is less unambiguous. THE FLORA OF RAUENBERG IN THE CONTEXT OF THE CONCEPT OF FLORISTIC COMPLEXES The plant-bearing marine sediments (Bodenheim Formation, Hochberg Member) are well correlated biostratigraphically by nannoplankton, dinolagellate cysts and the mammal genus Apterodon (Figure 1, Maxwell et al. 2016). The age is therefore limited Unangemeldet Heruntergeladen am | 10.04.17 15:37 384 to the middle Rupelian, i.e. approximately >30 Ma and < 33 Ma The stratigraphic constraints for the lora of Flörsheim are very similar (Kvaček 2004a, see also Introduction). The assemblages from Rauenberg and Flörsheim share many more taxa than any of the other compared loras. The Floristic Complex Nerchau-Flörsheim, irst established by Mai (1995), was further characterised by Kvaček and Walther (2001), who included also the assemblages of Markvartice and Suletice-Berand. The radiometric constraints of Suletice-Berand between 28.3 ± 1.4 Ma and 19.8 ± 0.5 Ma (Bellon et al. 1998) unambiguously indicate that loristically similar assemblages are not necessarily time equivalent. Assuming the validity of the radiometric dating, Suletice-Berand needs to be excluded. The age of the assemblage from Markvartice remains somewhat ambiguous (see above) and that of Nerchau is open (Mai & Walther 1991). Therefore, the typiication and deinition of the respective loristic complex should exclude Nerchau. Rauenberg is to be deinitely assigned to this complex. Due to the excellent stratigraphic correlation possibilities and the partly identical as well as complementary spectrum of taxa at Rauenberg, the Floristic Complex Flörsheim-Rauenberg is suggested here instead. Typiication for Rauenberg: Stratum typicum: Bodenheim Formation, Hochberg Member, “Fischschiefer”, NP 23, Lower Oligocene. Locus typicus: Rauenberg, Baden-Württemberg, Germany. The concept of loristic complexes presented by Kvaček and Walther (2001) is based on a north-south gradient (Boreal Province and Paratethys Province). The lora from the Tard Clay Formation serves as the type lora of the Floristic Complex Kiscell (Mai 1995, Kvaček & Walther 2001) at the southern margin of former Europe. The stratigraphic position and similarities of the lora from the Tard Clay Formation and Rauenberg (as well as Flörsheim) regarding joint taxa are clear including taxa which are not documented from the early Oligocene of North Bohemia or adjacent parts of Germany (Ceratozamia loersheimensis, Doliostrobus taxiformis, Sloanea olmediaefolia). This implies a more complex vegetation pattern in former Europe, one in which the regional climate was inluenced by marine straits, rather than a simple northsouth gradient. J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 ACKNOWLEDGEMENTS I am very grateful to Dr. M. Barbacka (Budapest), Prof. H. Freitag (Kassel), Prof. L. Hably, Prof. G. Kadereit, Dr. L. Kunzmann (Dresden), Prof. Z. Kvaček (Prag), Dr. F. Stauffer (Geneva), PD Dr. M. Thiv (Stuttgart), and Prof. A. Weber (Vienna) for scientiic discussions. Dr. G. Gregor (Munich), Dr. M. Rummel, J. Hendriks (Augsburg) and Dr. E. Frey and C. Eck (Karlsruhe ) enabled me to study the collections of Naturmuseum Augsburg and Staatliches Museum für Naturkunde Karlsruhe. Prof. Z. Kvaček (Prag), Dr. L. Kunzmann and C. Kunzmann (Dresden), and PD Dr. Wilde and K. Schmidt (Frankfurt/Main) provided the cuticular slides from different Bohemian sites and localities in Saxony and adjacent regions, and from Flörsheim for comparison. Dr. F. Stauffer provided modern leaf material of Berchemia from the Herbarium of the Jardin Botanique Genève. Dr. T. Denk (Stockholm) provided photographic material of cuticles from modern Illicium. R. Noll provided information on fossil material from the Alzey Formation. M. Kamenz (Stuttgart) patiently prepared all the cuticle material, K. Wolf-Schwenninger (Stuttgart) prepared the SEM images, and my husband H. Sommer (Stuttgart) provided the macro-photos and prepared the plates. I sincerely thank all of my colleagues and my husband, as well as Z. Kvaček and L. 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Lygodium kaulfussii Heer emend. Gardner et Ettingshausen 5. SMNS P 1952/55 6. SMNS P 1952/322 7. SMNS P 1952/332 8. Ceratozamia loersheimensis (Engelhardt) Kvaček, SMNS P P1952/456 9. Doliostobus taxiformis (Sternberg) Kvaček ? taxiformis Mai & Walther, NMA 2006-104/1497 10. Sequoia abietina (Brongniart in Cuvier) Knobloch, NMA 2016-3/2179 11–13. Taxodium sp. 11. SMNS P 1952/88 12. SMNS P 1953/103 13. SMNK 7710 14–15. Tetraclinis salicornioides (Unger) Kvaček 14. SMNS P 1952/118 15. SMNS P 1953/109 16. Pinus (Strobus) sp. 1, SMNS P 1952/292 17. Pinus (Strobus) sp. 2, SMNS P 1952/90 18. Pinus sp. 3, SMNS P 1952/460 19. Pinus sp. 4, SMNS P 1952/143 20. Pinus sp. 6, NMA 2006-111/1497 Scale bar: 10 mm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 1 J. Kovar-Eder Acta Palaeobot. 56(2) 391 Unangemeldet Heruntergeladen am | 10.04.17 15:37 392 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 2 1–6. Daphnogene cinnamomifolia (Brongniart) Unger 1. SMNS P 1952/71 2. SMNS P 1953/12 3. SMNS P 1953/8 4. SMNS P 1953/18 5. SMNS P1952/472 6. SMNS P 1953/76 7–8. Laurophyllum kinkelinii (Engelhardt) Kvaček 7. SMNS P1953/64 8. SMNS P 1952/184 9–11. Laurophyllum cf. kinkelinii (Engelhardt) Kvaček 2004 var. glabrum Kvaček 9. SMNS P 1952/396 10. SMNS P 1952/378 11. SMNS P 1953/87 12–16. Laurophyllum pseudoprinceps Weyland & Kilpper s.l. 12. SMNS P 1952/344 13. SMNS P 1953/48 14. SMNS P 1952/345 15. SMNS P 1953/84 16. SMNS P 1953/15A 17–23. Laurophyllum rauenbergense Kovar-Eder sp. nov. 17. SMNS P 1953/73 paratype 18. SMNS P 1952/350 19. SMNS P 1952/93 paratype 20. SMNS P 1953/82 holotype 21. SMNS P 1953/85 paratype 22. SMNS P 1952/379 23. SMNS P 1953/47 paratype 24–25. Laurophyllum cf. rauenbergense Kovar-Eder sp. nov. 24. SMNS P1952/231 25. SMNS P 1953/59 26. Actinodaphne pseudogermari Walther, SMNS P 1952/17 Scale bar: 10 mm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 2 J. Kovar-Eder Acta Palaeobot. 56(2) 393 Unangemeldet Heruntergeladen am | 10.04.17 15:37 394 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 3 1–2. Palmacites lamanonis Brongniart 1. SMNS P 1952/175 2. SMNS P 1953/124 3–4. Sabalites major (Unger) Saporta 3. SMNS P 1953/132 4. SMNS P 1952/169 5. Arecaceae gen. et sp. indet., NMA 2016-2/1497, scale 50 mm 6. ? Calamoideae gen. et sp. indet., NMA 2016-1/1497 7–9. Phoenicites sp. 7. SMNS P 1952/12 8. SMNS P 1952/141 9. NMA 2006-112/1497 10. Pinus engelhardtii Menzel, SMNK 7706 11–12. Pinus (Pinus) cf. thomasiana (Goeppert) Reichenbach 11. SMNS P 1952/176 12. SMNS P 1953/106 13. Pinus sp., SMNS P 1952/150 If not noted otherwise, scale bar: 10 mm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 3 J. Kovar-Eder Acta Palaeobot. 56(2) 395 Unangemeldet Heruntergeladen am | 10.04.17 15:37 396 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 4 1–10. Platanus neptuni (Ettingshausen) Bůžek, Holý et Kvaček 1. SMNS P 1952/32 2. SMNS P 1952/81 3. SMNS P 1952/14 4. SMNS P 1952/26 5. SMNS P 1952/220 6. SMNS P 1952/86 7. NMA 2006-128/1497, 7a. venation details, scale bar 5 mm 8. SMNS P 1952/37 9. SMNS P 1952/24A 10. SMNS P 1952/19 11. Myrica cf. lignitum (Unger) Saporta sensu Ettingshausen & Standfest, SMNS P 1952/361 12–17. Myrica obliquifolia sp. nov. 12. SMNS P 1952/238 paratype 13. SMNS P 1952/232 paratype 14. SMNS P 1952/162 paratype 15. SMNS P 1952/255 paratype 16. SMNS P 1953/99 paratype 17. SMNS P 1952/85 holotype 18–20. Myrica longifolia Unger 18. NMA 2006-105/1497 19. SMNS P 1952/50 20. SMNS P 1952/186 If not noted otherwise, scale bar: 10 mm Unangemeldet Heruntergeladen am | 10.04.17 15:37 397 Plate 4 J. Kovar-Eder Acta Palaeobot. 56(2) Unangemeldet Heruntergeladen am | 10.04.17 15:37 398 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 5 1. Phoenicites sp., NMA 2006-180/1497, scale bar 100 mm 2–3. Sloanea olmediaefolia (Unger) Z. Kvaček & Hably 2. NMA 2006-114/1497, 2a. venation details 3. SMNS P 1952/340 4. Sloanea artocarpites (Ettingshausen) Z. Kvaček et Hably, P1952/61 5. ? Berchemia altorhenana Kovar-Eder sp. nov., SMNS P 1953/92, holotype 6. Populus germanica (Menzel) Walther, SMNK 7568 7. ? Ternstroemites maritiae Kovar-Eder sp. nov., SMNS P 1952/342, holotype 8–10. Symplocos deichmuelleri (Kvaček & Walther) comb. nov. 8. SMNS P1952/62 9. SMNS P1952/65 10. SMNS P 1953/43 11. Pungiphyllum cruciatum (Al. Braun) Frankenhäuser & Wilde, NMA 2006-126/1497 12–14. Oleinites rauenbergensis Kovar-Eder sp. nov. 12. SMNS P 1952/299 paratype 13. SMNS P 1952/402 paratype 14. SMNS P 1952/380 holotype 15–16. ? Oleinites rauenbergensis Kovar-Eder sp. nov. 15. SMNS P 1952/6 16. SMNS P 1953/93 17–18. Oleinites altorhenana Kovar-Eder sp. nov. 17. SMNS P 1952/191 holotype 18. SMNS P 1953/62 paratype If not noted otherwise, scale bar: 10 mm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 5 J. Kovar-Eder Acta Palaeobot. 56(2) 399 Unangemeldet Heruntergeladen am | 10.04.17 15:37 400 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 6 1. Betula dryadum Brongniart, SMNK 7711, scale bar 1 mm 2. Craigia bronnii (Unger) Kvaček, Bůžek et Manchester, SMNK 7748 3. Winged fruit or seed incertae sedis, NMA 2006-113/1497 4. Hydrangea microcalyx Sieber, SMNS P 1952/145 5. ? Leguminocarpon sp., NMA 2006-173/1497 6–7. Comptonia difformis (Sternberg) Berry 6. SMNS P 1953/118 7. SMNS P 1952/166 8. Laurophyllum sp. A, SMNS P 1952/336 9. Laurophyllum sp. B, SMNS P 1952/47 10. Laurophyllum sp. C, SMNS P 1953/81 11–13. Laurus abchasica (Kolakovskii & Shakryl) Ferguson 11. SMNS P 1952/382 12. SMNS P 1953/75 13. SMNS P 1953/65 14. Eotrigonobalanus furcinervis (Rossmässler) Walther & Kvaček forma haselbachensis (Kvaček & Walther) Kvaček & Walther in Walther, SMNS P 1953/60 15–17. Engelhardia orsbergensis (Wessel & Weber) Jähnichen, Mai & Walther 15. NMA 2006-108/1497 16. SMNK 7499 17. SMNS P 2228/1 18. Engelhardia macroptera (Brongniart) Unger, SMNS P 1952/154 19. ? Buxus egeriana Kvaček, Bůžek et Holý, NMA 2006-168/1497 20. Betulaceae vel Ulmaceae gen. et sp. indet., SMNS P 1952/437 21. Carya quadrangula (Kirchheimer) Leroy, SMNS P 1952/159 22. Carya fragiliformis (Sternberg) Kvaček & Walther, SMNS P 1952/58 23. Distylium metzleri Kovar-Eder sp. nov., SMNS P 1952/43 holotype 24–25. Trachelospermum kelleri Kovar-Eder sp. nov. 24. SMNS P 1953/101 paratype 25. NMA 2006–121/1497 holotype 26–27. Trachelospermum steiningeri Kvaček 26. SMNS P 1952/101 27. SMNS P 1952/366 If not noted otherwise, scale bar: 10 mm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 6 J. Kovar-Eder Acta Palaeobot. 56(2) 401 Unangemeldet Heruntergeladen am | 10.04.17 15:37 402 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 7 1–5. Viscophyllum hendriksiae Kovar-Eder sp. nov. 1. SMNS P 1952/11 paratype 2. SMNS P 1952/57 paratype 3. SMNS P 1953/70 holotype 4. SMNS P 1953/77 paratype 5. SMNS P 1952/158 paratype 6. Dicotylophyllum ziegleri Kovar-Eder sp. nov., SMNS P 1953/86 holotype 7–8. Dicotylophyllum vesiculaeferens Kovar-Eder sp. nov. 7. SMNS P 1952/310 holotype 8. SMNS P 1952/306 paratype 9–14. Cladites vesiculaeferens Kovar-Eder sp. nov. 9. SMNS P 1952/371 paratype 10. SMNS P 1952/416 holotype 11. NMA 2006-154/1497 12. SMNS P 1953/105 13. NMA 2016-4/2179 14. SMNS P 1952/87 paratype 15. Spiny branch, NMA 2006-23/1497 16. Dicotylophyllum badense Kovar-Eder sp. nov., SMNS P 1952/200 holotype 17. Dicotylophyllum oechsleri Kovar-Eder sp. nov., SMNS P 1952/454 holotype 18. Dicotylophyllum sp., SMNS P 1952/400 19. Cladites vesiculaeferens Kovar-Eder sp. nov., SMNS P 1952/104 Scale bar: 10 mm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 7 J. Kovar-Eder Acta Palaeobot. 56(2) 403 Unangemeldet Heruntergeladen am | 10.04.17 15:37 404 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 8 1–3. Microthyriaceae gen. et sp. indet. Different stages of fruiting bodies on the cuticle of Phoenicites sp. 1. SMNS P 1952/473-1 2. SMNS P 1952/473-2 3. SMNS P 1952/473-1 4–7. Ceratozamia loersheimensis (Engelhardt) Kvaček 4–6. Cuticle in stoma-bearing condition, note the short rows of strongly cutinised short cells; 4. SMNS P 1952/456-2 5. SMNS P 1952/456-1 6. Stomata with encircling cells, polar cells very slender, remnants of guard cells still preserved, SMNS P 1952/456-1 7. Cuticle in non-stoma-bearing condition, SMNS P 1952/456-2 8. Sequoia abietina (Brongniart in Cuvier) Knobloch, NMA 2016-3/2179-2 9. Tetraclinis salicornioides (Unger) Kvaček, SMNS P 1952/194-2 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 8 J. Kovar-Eder Acta Palaeobot. 56(2) 405 Unangemeldet Heruntergeladen am | 10.04.17 15:37 406 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 9 1–3. Actinodaphne pseudogermari Walther 1. Adaxial cuticle and oil glands, SMNS P 1952/17-1 2–3. Abaxial cuticle with stomata overlapped by neighbouring cells, SMNS P 1952/17-2 4–9. Daphnogene cinnamomifolia (Brongniart) Unger 4–5. Probable shade leaf 4. Adaxial cuticle with undulate anticlines and a single trichome base upon a lower-order vein, SMNS P 1953/8-2 5. Abaxial cuticle, largely glabrous except for a prominent trichome base, SMNS P 1953/8-1 6. Abaxial cuticle, trichome bases with thickened polar rim, SMNS P 1953/76-2 7–8. Probable sun leaf 7. Adaxial cuticle with rather straight anticlines, SMNS P 1952/76-2 8. Abaxial cuticle with trichome bases, SMNS P 1952/76-2 9. Abaxial cuticle of a probable sun leaf with dense trichome bases, SMNS P 1952/472-1 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 407 Plate 9 J. Kovar-Eder Acta Palaeobot. 56(2) Unangemeldet Heruntergeladen am | 10.04.17 15:37 408 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 10 1–3. Laurus abchasica (Kolakovskii & Shakryl) Ferguson 1. Adaxial cuticle with ? hypodermis, SMNS P 1953/65-2 2–3. Abaxial cuticle 2. SMNS P 1953/65-2 3. SMNS P 1953/75-2 4–9. Laurophyllum rauenbergense Kovar-Eder sp. nov. 4. Adaxial cuticle, SMNS P1953/82-1, holotype 5. Abaxial cuticle, non-modiied cells somewhat domed, stomata indistinct, SMNS P 1953/82-2, holotype 6. Abaxial cuticle, non-modiied cells somewhat domed, stomata indistinct, SMNS P 1952/391-1, paratype 7. Adaxial cuticle with distinct, large oil glands, SMNS P 1952/93-1, paratype 8–9. Abaxial cuticle 8. SMNS P 1952/445-2, paratype 9. SMNS P 1953/47-1, paratype Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 10 J. Kovar-Eder Acta Palaeobot. 56(2) 409 Unangemeldet Heruntergeladen am | 10.04.17 15:37 410 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 11 1–4. Laurophyllum ? rauenbergense Kovar-Eder sp. nov. 1. Adaxial cuticle with slight striae around trichome base, SMNS P 1952/231-2 2. Abaxial cuticle SMNS P 1952/231-1 3. Adaxial cuticle SMNS P 1953/59-1 4. Abaxial cuticle SMNS P 1953/59-1 5–9. Laurophyllum kinkelinii (Engelhardt) Kvaček 5. Adaxial cuticle, SMNS P 1952/184-1 6–7. Abaxial cuticle with distinctly domed non-modifed cells, SMNS P 1952/184-3 7. Close-up view of 6. 8. Adaxial cuticle, SMNS P 1953/64-1 9. Abaxial cuticle, SMNS P 1953/64-2 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 11 J. Kovar-Eder Acta Palaeobot. 56(2) 411 Unangemeldet Heruntergeladen am | 10.04.17 15:37 412 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 12 1–4. Laurophyllum cf. kinkelinii (Engelhardt) Kvaček 2004 var. glabrum Kvaček 1. Adaxial cuticle, SMNS P 1952/378-1 2. Abaxial cuticle, trichome bases large with radial striae, SMNS P 1952/378-2 3. Adaxial cuticle with striae upon veins and trichome base, SMNS P 1953/87-1 4. Abaxial cuticle 4a. Tichome bases, SMNS P 1953/87-1 4b. Domed cells and sunken, overlapped stomata, SMNS P 1953/87-2 5–7. Laurophyllum pseudoprinceps Weyland & Kilpper s.l. 5. Adaxial cuticle, SMNS P 1952/389-2 6. Abaxial cuticle, SMNS P 1952/389-2 7. Abaxial cuticle, SMNS P 1953/15A-2 8–9. Laurophyllum sp. A 8. Adaxial cuticle, SMNS P 1952/336-2 9. Abaxial cuticle, SMNS P 1952/336-1 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 12 J. Kovar-Eder Acta Palaeobot. 56(2) 413 Unangemeldet Heruntergeladen am | 10.04.17 15:37 414 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 13 1–3. Laurophyllum sp. B 1. Adaxial cuticle with a single oil gland, SMNS P 1952/47-1 2–3. Abaxial cuticle, SMNS P 1952/47-1 3. Close-up view of 2. 4. Laurophyllum sp. C 4a. Adaxial cuticle, SMNS P 1953/81-2 4b. Abaxial cuticle, SMNS P 1953/81-2 5. Palmacites lamanonis Brongniart, cuticle in stoma-bearing condition, loosely spaced brachyparatetracytic stomatal complexes, 1952/66-1 6–7a. ? Calamoideae gen. et sp. indet., cuticle in stoma-bearing condition, paratetracytic to cyclocytic stomatal complexes, NMA 2016-1/1497-1 6. Close-up view of 7a. 7b–8. Arecaceae gen. et sp. indet., cuticle in stoma-bearing condition, NMA 2016-2/1497-1 8. Close-up view of 7b. 9. Phoenicites sp., Cuticle in stoma-bearing condition, SMNS P 1952/12-1 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 13 J. Kovar-Eder Acta Palaeobot. 56(2) 415 Unangemeldet Heruntergeladen am | 10.04.17 15:37 416 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 14 1–6. Platanus neptuni (Ettingshausen) Bůžek, Holý et Kvaček 1. Adaxial cuticle, minutely undulate anticlines and a single gland with at least three base cells, SMNS P 1952/32-1 2. Abaxial cuticle, anticlines vaguely visible, numerous glands, SMNS P1952/32-1 3. Abaxial cuticle, distinct striae to wrinkles, SMNS P 1952/37-1 4. Adaxial cuticle, undulation coarser than in 1., several glands and striation upon veins, SMNS P 1952/465-2 5. Abaxial cuticle, stomata in subparallel position along the margin, SMNS P 1952/465-2 6. Abaxial cuticle, stomatal ledges extremely pronounced but anticlines hardly visible, SMNS P 1953/44-1 7. Eotrigonobalanus furcinervis (Rossmässler) Walther & Kvaček forma haselbachensis (Kvaček & Walther) Kvaček & Walther in Walther, abaxial cuticle, roundish stomatal complexes and simple trichome bases, SMNS P 1953/60-1 8. Betulaceae vel Ulmaceae gen. et sp. indet., abaxial cuticle, SMNS P 1952/437-1 9. Myrica cf. lignitum (Unger) Saporta sensu Ettingshausen & Standfest 9a. Abaxial cuticle with stomata, SMNS P 1952/361-1 9b. Abaxial cuticle with stoma and peltate trichome, SMNS P 1952/361-2 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 417 Plate 14 J. Kovar-Eder Acta Palaeobot. 56(2) Unangemeldet Heruntergeladen am | 10.04.17 15:37 418 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 15 1–2. Myrica longifolia Unger 1. Abaxial cuticle, stoma (arrow), four-celled trichome base, SMNS P 1952/50-1 2a. ? Adaxial cuticle, SMNS P 1952/186-1 2b. Abaxial cuticle, pluricellular trichome bases, SMNS P 1952/186-1 3–9. Myrica obliquifolia sp. nov. 3a. Adaxial cuticle, SMNS P 1952/85-1, holotype 3b. Peltate trichome with oblong, two-celled trichome base, SMNS P 1952/85-1, holotype 4. Adaxial cuticle, SMNS P 1952/232-4, paratype 5a–b. Peltate trichomes with oblong, two-celled trichome bases 5a. SMNS P 1952/232-5 paratype 5b. SMNS P 1952/232-4 paratype 6. Abaxial cuticle, mainly stoma space visible, except for a single one preserved (arrow), SMNS P 1952/238-1, paratype 7. Abaxial cuticle, stomata, SMNS P 1952/255-1, paratype 8. Adaxial cuticle, SMNS P 1952/255-2, paratype 9a–b. Peltate trichome focused differently, SMNS P 1953/99-2, paratype 9a. Focus on trichome base 9b. Focus on peltate trichome Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 15 J. Kovar-Eder Acta Palaeobot. 56(2) 419 Unangemeldet Heruntergeladen am | 10.04.17 15:37 420 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 16 1–3. Distylium metzleri Kovar-Eder sp. nov. 1. Abaxial cuticle, concentric striae upon stomata, parallel wrinkles upon veins and parallel to radial ones around trichome base, SMNS P 1952/43-1, holotype 2. Abaxial cuticle, concentric striae upon stomata, SMNS P 1952/43-2, holotype 3. Adaxial cuticle, SMNS P 1952/43-2, holotype 4–6. Sloanea olmediaefolia (Unger) Z. Kvaček & Hably 4a. Adaxial cuticle, NMA 2006-114/1497-2 4b. Abaxial cuticle, stomata roundish, cyclo- to anomocytic, NMA 2006-114/1497-2 5. Abaxial cuticle, stomata roundish, cyclo- to anomocytic, SMNS P 1952/340-1 6. Adaxial cuticle, SMNS P 1952/340-1 7–9. ? Berchemia altorhenana Kovar-Eder sp. nov. 7. Adaxial cuticle, zig-zag coarse of anticlines, SMNS P 1953/92-2, holotype 8. Abaxial cuticle, stomata masked by hyphae of fungi (arrow), trichome bases, SMNS P 1953/92-1, holotype 9. Abaxial cuticle, several stomata, SMNS P 1953/92-2, holotype Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 16 J. Kovar-Eder Acta Palaeobot. 56(2) 421 Unangemeldet Heruntergeladen am | 10.04.17 15:37 422 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 17 1–3. ? Ternstroemites maritiae Kovar-Eder sp. nov. 1a. Adaxial cuticle with strongly undulate anticlines, SMNS P 1952/342-1, holotype, 1b. Abaxial cuticle, SMNS P 1952/342-1, holotype 2–3. Abaxial cuticle, SMNS P 1952/342-2, holotype 4–6. Symplocos deichmuelleri (Kvaček & Walther) comb. nov. 4. Adaxial cuticle, SMNS P 1952/62-1 5–6. Abaxial cuticle, SMNS P 1952/65-1 7–9. Trachelospermum steiningeri Kvaček 7. Adaxial cuticle with remains of secretory bodies, SMNS P 1952/366-2 8–9. Abaxial cuticle, stomata cyclocytic, remains of secretory bodies, SMNS P 1952/366-2 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 17 J. Kovar-Eder Acta Palaeobot. 56(2) 423 Unangemeldet Heruntergeladen am | 10.04.17 15:37 424 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 18 1–2. Trachelospermum steiningeri Kvaček 1. Adaxial cuticle with numerous, well-preserved secretory bodies, SMNS P 1952/101-2 2. Abaxial cuticle, stomata (arrows), secretory bodies, SMNS P 1952/101-2 3–5. Trachelospermum kelleri Kovar-Eder sp. nov. 3. Abaxial cuticle, secretory bodies dense, masking the stomata (arrows), NMA 2006-121/1497-1, holotype 4. Abaxial cuticle, a single stoma and its aperture visible, NMA 2006-121/1497-1, holotype 5. Abaxial cuticle, secretory bodies masking the stomata (arrows), SMNS P 1953/101-3, paratype 6–9. Oleinites altorhenana Kovar-Eder sp. nov. 6. Adaxial cuticle, SMNS P 1952/191-2, holotype 7. Abaxial cuticle, trichome bases, SMNS P 1952/191-2, holotype 8a. Abaxial cuticle, trichome base and small non-modiied epidermal cells, SMNS P 1952/191-2, holotype 8b. Abaxial cuticle, stomata, SMNS P 1952/191-2, holotype 9. Abaxial cuticle, stomata and a single trichome base, SMNS P 1952/388-2, paratype Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 18 J. Kovar-Eder Acta Palaeobot. 56(2) 425 Unangemeldet Heruntergeladen am | 10.04.17 15:37 426 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 19 1–5a. Oleinites rauenbergensis Kovar-Eder sp. nov. 1. Adaxial cuticle, SMNS P 1952/380-2, holotype 2a. Probable abaxial cuticle, a prominent trichome base with small peltate head, SMNS P 1952/380-2, holotype 2b. Abaxial cuticle, stomata, SMNS P 1952/380-2, holotype 3a–c. Small fragments of abaxial cuticle with single stomata 3a–b. SMNS P 1952/380-2, holotype 3c. SMNS P 1952/402-2, paratype 4. Adaxial cuticle, SMNS P 1952/402-2, paratype 5a. Abaxial cuticle, SMNS P 1952/402-1, paratype 5b–6. ? Oleinites rauenbergensis Kovar-Eder sp. nov. 5b. Cuticle fragment with trichome bases, SMNS P 1952/6-1 6. Abaxial cuticle, SMNS P 1952/6-1 7–9. Dicotylophyllum badense Kovar-Eder sp. nov. 7. Adaxial cuticle, SMNS P 1952/200-2, holotype 8. Abaxial cuticle, SMNS P 1952/200-3, holotype 9a–b. Abaxial cuticle, stomata 9a. SMNS P 1952/200-2, holotype 9b. SMNS P 1952/200-3, holotype Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 427 Plate 19 J. Kovar-Eder Acta Palaeobot. 56(2) Unangemeldet Heruntergeladen am | 10.04.17 15:37 428 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 20 1–5. Dicotylophyllum vesiculaeferens Kovar-Eder sp. nov. 1. Adaxial cuticle, SMNS P 1952/310-1, holotype 2. Abaxial cuticle with a group of penta- to hexagonal pouches, some still with secretory content, SMNS P 1952/310-1, holotype 3. Close-up view showing balloon-shaped secretory structures in pouches and empty pouches each with a plurilet central mark (arrow), SMNS P 1952/310-1, holotype 4. Abaxial surface, ? stomata (arrows) and single pouches, SMNS P 1952/310-1, holotype 5. ? Abaxial cuticle with a illed pouch, SMNS P 1952/310-2, holotype 6–9. Cladites vesiculaeferens Kovar-Eder sp. nov. 6. Cuticle of the axis, cells arranged in rows and pouches, SMNS P 1952/416-1, holotype 7a–b. and 8a–b. Pouches in different close-up views with adherent and embedded balloon-shaped secretory bodies, SMNS P 1952/416-1, holotype 9. Cuticle from a more proximal part of the same axis, cuticle more delicate, cell arrangement not visible, pouches in different stages of development, SMNS P 1952/416-2, holotype Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 20 J. Kovar-Eder Acta Palaeobot. 56(2) 429 Unangemeldet Heruntergeladen am | 10.04.17 15:37 430 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 21 1–3. Dicotylophyllum vesiculaeferens Kovar-Eder sp. nov., SEM view of SMNS P 1952/310, holotype 1–2. External view of secreting structure 3. Internal view of secreting structure 4–8. Cladites vesiculaeferens Kovar-Eder sp. nov. 4–6. Internal view, partly clustered secreting structures, i.e. pouches partly with still adherent vesicles, SEM view of SMNS P 1952/416, holotype 4. scale bar 50 µm 6. vesicles appear interconnected 7, 8. External view, SMNS P 1952/104 9. Dicotylophyllum ziegleri Kovar-Eder sp. nov., SEM view of abaxial cuticle, internal side, two cyclocytic stomata and a single gland, SMNS P 1953/86, holotype, scale bar 50 µm If not noted otherwise, scale bar: 5 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 21 J. Kovar-Eder Acta Palaeobot. 56(2) 431 Unangemeldet Heruntergeladen am | 10.04.17 15:37 432 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 22 1–4. Dicotylophyllum oechsleri Kovar-Eder sp. nov. 1. Adaxial cuticle, strongly sinuous anticlines, distinct striation, base of a glandular trichome SMNS P 1952/454-1, holotype 2a. Adaxial cuticle, SMNS P 1952/454-2, holotype 2b–4b. Abaxial cuticle, SMNS P 1952/454-1, holotype 2b. Stoma and mesophyll remains 3. Stomata and prominent trichome base of a disc-shaped or globular head 4a–b. Trichome base with depression of the head 4a. Focus on the marginal ridge of the head 4b. Same position as 4a. but focus on the collar-like margin of the trichome base 5–7. Dicotylophyllum ziegleri Kovar-Eder sp. nov. 5a. Adaxial cuticle, SMNS P1953/86-1, holotype 5b. Abaxial cuticle, subparallel stoma orientation, stomata cyclo- to actinocytic, somewhat grouped, SMNS P 1953/86-2, holotype 6a–b. Epidermal glands marginally overlapped by the surrounding, marginally strongly thickened cells 6a. SMNS P 1953/86-2, holotype 6b. SMNS 1953/86-1, holotype 7. Abaxial cuticle with stomata and gland, SMNS P 1953/86-2, holotype 8–9. Dicotylophyllum sp. 8a–b. Abaxial cuticle, SMNS P 1952/400-1 8a. Cell outlines more or less vaguely visible, ? stomata, trichome bases 8b. A single ? stoma (arrow) 9. Adaxial cuticle with large trichome bases, SMNS P 1952/400-1 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 22 J. Kovar-Eder Acta Palaeobot. 56(2) 433 Unangemeldet Heruntergeladen am | 10.04.17 15:37 434 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Plate 23 1–6. Viscophyllum hendriksiae Kovar-Eder sp. nov. 1. Adaxial cuticle, SMNS P 1953/70-1, holotype 2. Abaxial cuticle, cells more or less arranged in rows, subparallel stoma orientation, SMNS P 1953/70-1, holotype 3. Abaxial cuticle, close-up, SMNS P 1953/70-1, holotype 4. Abaxial cuticle, SMNS P 1952/11-1, paratype 5. Adaxial cuticle, ? hypodermis, SMNS P 1953/71-2, paratype 6. Abaxial cuticle, SMNS P 1953/71-2, paratype 7. Berchemia philippinensis Vid., abaxial cuticle, non-modiied cells domed, partly overlapping stomata, surface granular, SMNS 2015/18-1 8. Berchemia lineata (L.) DC., abaxial cuticle, stomata partly overlapped by non-modiied cells, SMNS 2015/16-1 9. Berchemia afinis Hassk., adaxial cuticle, anticlines of non-modiied cells somewhat zig-zag undulate, trichome base with intensively staining foot, SMNS 2015/7-1 Scale bar: 50 µm Unangemeldet Heruntergeladen am | 10.04.17 15:37 Plate 23 J. Kovar-Eder Acta Palaeobot. 56(2) 435 Unangemeldet Heruntergeladen am | 10.04.17 15:37 436 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Appendix 1. Cuticular features of ? Berchemia altorhenana Kovar-Eder sp. nov. compared to modern species of Berchemia Abaxial surface Berchemia species stomata stoma length (average) × width (average), aperture length (AL), average in brackets, [µm] 24–38 (35) 9–13 (11) × 6–11 ( 9), AL = 4–6 (5) somewhat shallowly undulate not domed 10–15 (12) not reproduced on cuticle, straight-bent hardly reproduced on cuticle, straight-bent hardly reproduced on cuticle, bent delicate, straight somewhat domed not measurable hardly reproduced on cuticle domed not measurable not domed 13–26 (20) strongly domed, papillae-like strongly domed not measurable not measurable 16–23 (20) B. berchemiaefolia Koidz. not to somewhat sunken B. chanetii Lév. not sunken 12–18 (14) × 9–12 (11), AL = 6–11 (9) B. edgeworthii Law. not sunken AL = 8–11 (9) B. loribunda (Wall.) Brongn. not sunken, AL = 10–14 (13) orientation partly subparallel sunken AL = 13–20 (15) B. fournieri Pancher not sunken B. giraldiana Schn. sunken B. kulingensis Schneid. B. lineata (L.) CD. sunken B. longeracemosa Okuyama 13–15 × 7–10, AL = 8–14 (10) 15–19 (16) × 11–14 (13), distinct, slender, AL = 8–11 (9) straight to somewhat wavy AL = 9–12 (11) not reproduced on cuticle AL = 8–11 (10) not reproduced on cuticle 13–18 (15) × ??, delicate, straight AL = 8–13 (10) sunken, orientation partly subparallel not sunken, 14–20 (16) × ??, partly subparallel AL=8–12 (10) B. philippinensis Vid. distinctly sunken hardly reproduced on cuticle, straight-bent AL = 9–12 (10) not reproduced on cuticle 13–17 (15) × 12–13 (12), delicate, straight to AL = 9–11 (10) bent, only reproduced along veins AL= 9–13 (11) not reproduced on cuticle B. racemosa S. & Z. distinctly sunken B. sinica Schn. not sunken, orientation partly subparallel distinctly sunken AL = 7–11 (9) B. yunnanensis Franch. size of nonmodiied epidermal cells, [µm] not domed B. afinis Hassk. B. lavescens (Wall.) Brongn. surface of non-modiied epidermal cells 17–25 (21) × 10–21 (15), moderately AL = 10–17 undulate slightly sunken, overlapped by surrounding cuticle not sunken B. altorhenana Kovar-Eder sp. nov. anticlines of non-modiied epidermal cells not reproduced on cuticle not domed not measurable somewhat domed not measurable not domed 23–45 (36) domed slightly domed not measurable strongly domed, papillae-like strongly domed, papillae-like not measurable not measurable slightly domed not measurable strongly domed, papillae-like 17–27 (22) Unangemeldet Heruntergeladen am | 10.04.17 15:37 437 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Appendix 1. Continued Adaxial cuticle trichomes rare granulate glabrous somewhat granulate surface size of nonanticlines of of nonmodiied non-modiied modiied indumentum epidermal epidermal cells epidermal cells, [µm] cells moderately zig-zag not domed 24–38 (35) scattered trichome bases, undulate dark-staining centre, sometimes ring of small encircling cells slender, smooth, not domed 15–31 (22) scattered trichome bases, moderately a bit zig-zag dark-staining center, undulate ring of small encircling cells hardly reproduced on slightly not trichome bases rare with cuticle, straight-bent domed measurable ring of short girdling cells hardly reproduced on the not domed not glabrous cuticle, straight-bent measurable glabrous granulate distinct, smooth, straight not domed 18–39 (27) glabrous granulate glabrous inely granulate largely glabrous roughly granulate glabrous inely granulate very thick, straight, con- not domed 27–51 (34) tinuous, resolving into pits to vaguely visible distinct, straight, smooth not domed 29–44 (36) glabrous glabrous rather smooth, occasionally delicate folding granulate indumentum trichome bases scattered, not dense absent trichomes abundant glabrous idiocuticular structure smooth smooth granulate distinct, slender, straight, smooth but pitted on the inner side hardly reproduced on cuticle, straight faintly reproduced on cuticle, straight vaguely visible, straight, slender idio-cuticular structure smooth smooth granulate granulate not domed 16–36 (24) glabrous not domed not measurable not domed 42–60 (53) glabrous inely granulate, additionally scattered folds somewhat inely granulate granulate glabrous granulate not domed 26–47 (33) glabrous granulate to rugulate 32–43 (37) glabrous granulate glabrous smooth glabrous granulate vaguely visible, straight, slightly slender domed glabrous strongly granulate strongly granulate thick, straight, smooth not domed 13–20 (17) glabrous granulate thick, straight, smooth, desintegrating into pits not domed 20–36 (30) glabrous granulate glabrous granulate thick, straight, smooth not domed 19–33 (26) glabrous somewhat granulate trichomes abundant smooth thick, straight, smooth, desintegrating into pits to hardly traceable not domed 17–27 (22) glabrous granulate glabrous Unangemeldet Heruntergeladen am | 10.04.17 15:37 438 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 Cystoseirites communis Thallites multiidus Laminarites latus Lygodium kaulfussii Filicopsida sp. Ceratozamia loersheimensis Doliostobus taxiformis ? taxiformis Sequoia abietina Taxodium sp. Tetraclinis salicornioides Pinus (Strobus) sp. 1 Pinus (Strobus) sp. 2 Pinus sp. 3 Pinus sp. 4 Pinus sp. 5 Pinus sp. 6 Pinus (Pinus) cf. thomasiana Pinus engelhardtii Actinodaphne pseudogermari Daphnogene cinnamomifolia Laurus abchasica Laurophyllum kinkelinii inkl. L. kinkelinii glabrum Laurophyllum pseudoprinceps Laurophyllum rauenbergense Laurophyllum sp. A Laurophyllum sp. B Laurophyllum sp. C Palmacites lamanonis Sabalites major Phoenicites sp. ? Calamoideae gen. et sp. indet. Arecaceae gen. et sp.indet. Platanus neptuni ? Buxus egeriana Eotrigonobalanus furcinervis f. haselbachensis Betula dryadum Betulaceae vel Ulmaceae gen. et sp. indet. Comptonia difformis Myrica cf. lignitum Myrica longifolia Myrica obliquifolia Carya fragiliformis Carya quadrangula Engelhardia orsbergensis/ E. macroptera ? Berchemia altorhenana ? Leguminocarpon sp. Sloanea artocarpites Sloanea olmediaefolia Populus germanica 1.0 1.0 1.0 0.5 0.5 0.5 0.5 1.0 0.5 0.5 0.5 0.5 1.0 1.0 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 1.0 0.5 0.5 1.0 0.5 1.0 0.5 1.0 0.5 0.5 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 Control aquatic azonal nonwoody azonal woody m-herb d-herb arboreal fern LEG SCL BLE BLD CONIF Taxon Azonal arboreal palm Zonal Problematic taxa/ uncounted Appendix 2. Score sheet of the IPR vegetation analysis (after Kovar-Eder & Kvaček 2008, Teodoridis et al. 2011a) 0.5 0.5 0.5 0.5 0.5 1.0 0.5 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 0.5 0.3 0.3 0.5 0.5 0.5 0.3 1.0 0.3 0.3 0.3 0.5 0.5 0.3 0.5 0.5 1.0 1.0 1.0 1.0 1.0 1.0 0.3 1.0 0.5 1.0 1.0 1.0 1.0 1.0 0.3 1.0 0.5 0.5 0.5 0.5 0.5 Unangemeldet Heruntergeladen am | 10.04.17 15:37 439 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 sum of taxa sum of zonal taxa percent of all zonal taxa 0.5 0.5 aquatic azonal nonwoody azonal woody m-herb d-herb arboreal fern LEG SCL BLE Control Craigia bronnii Distylium metzleri ? Viscophyllum hendriksiae Hydrangea microcalyx ? Ternstroemites maritiae Symplocos deichmuelleri Trachelospermum kelleri Trachelospermum steiningeri Oleinites altorhenana Oleinites rauenbergensis Pungiphyllum cruciatum Dicotylophyllum badense Dicotylophyllum oechsleri Dicotylophyllum vesiculaeferens Dicotylophyllum ziegleri Dicotylophyllum sp. Cladites vesiculaeferens Winged fruit or seed incertae sedis Spiny branch BLD CONIF Taxon Azonal arboreal palm Zonal Problematic taxa/ uncounted Appendix 2. Continued 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 1.0 0.5 0.5 1.0 1.0 0.3 22.8 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.3 4.8 1.5 2.5 0.0 0.0 1.0 11.7 18.8 48.6 10.3 3.2 5.3 0.0 0.0 2.1 5.5 8.8 1.0 1.0 0.5 0.5 1.0 0.5 0.5 0.5 0.5 0.5 0.5 0.3 14.0 1.0 3.0 3.0 68.0 47.0 100.0 Unangemeldet Heruntergeladen am | 10.04.17 15:37 MAT – mean annual temperature, CMMT – coldest month mean temperature, WMMT – warmest month mean temperature, MAP – mean annual precipitation, WMMP – wettest month mean precipitation, DMMP – driest month mean precipitation. Climate data derived from http://de.climate-data.org Warmest month WMMT [°C] Coldest month CMMT [°C] MAP [mm] Wettest month WMMP [mm] Driest month DMMP [mm] Climate type sensu Köppen 29 397 28.2 24.6 April May 31.3 27.6 Dec Jan 24.7 19.5 1143 1737 Sept Aug 249 434 Dec Feb 4 8 Aw Aw 16°04′N 21°02′N 22°29′N 10 21 112 25.7 23.8 23.8 June June June 29.2 29.4 28.3 Jan Jan Jan 21.4 16.6 17.1 2075 1684 1735 Oct Aug Aug 553 338 330 March Jan Jan 30 16 18 Am Cwa Cwa 110°15′E 107°22′E 108°13′E 109°37′E 113°29′E 116°41′E 102°41′E 110°17′E 114°56′E 20°00′N 22°25′N 22°38′N 23°06′N 23°13′N 23°24′N 25°04′N 25°16′N 25°50′N 8 106 81 52 18 15 1911 155 120 24.2 22.8 22.5 21.9 22.2 21.7 15.2 19.5 19.8 July July July July July July July July July 28.9 29.0 29.1 29.0 28.8 28.3 20.3 28.7 29.8 Jan Jan Jan Jan Jan Feb Jan Jan Jan 17.5 14.5 13.7 12.9 13.9 14.2 8.4 9.0 8.4 1581 1518 1363 1469 1720 1527 999 1775 1467 Sept June June May June June July May June 264 270 228 258 287 278 209 347 232 Jan Dec Dec Nov Dec Dec Jan Dec Dec 25 30 30 38 32 31 12 48 39 Cwa Cfa Cfa Cfa Cfa Cfa Cwb Cfa Cfa Mexico 15 Guanajuato, Gto 16 Veracruz 101°17′W 96°08′W 21°00′N 19°09′N 2060 8 17.6 25.4 May June 20.8 28.1 Jan Jan 13.9 21.7 697 1516 July July 145 360 Feb Feb 7 13 Cwb Aw Guatemala 17 Aguacatan 18 Puerta Parada 20 San Raymundo 19 Olopa 91°18′W 90°27′W 90°36′W 89°20′W 15°20′N 14°33′N 14°46′N 14°41′N 3093 1846 1578 1325 10.2 17.2 18.6 19.9 May April May April 11.2 18.4 19.8 21.7 Jan Jan Jan Dec 8.6 15.7 17.0 17.7 1457 1529 1578 1526 Sept June June June 258 315 225 284 Feb Feb Feb Jan 19 5 2 28 Cwb Cwb Cwb Cwb Latitude Thailand 1 Nakhon Sawan 2 Chiang Rai 100°08′E 99°50′E 15°42′N 19°55′N Vietnam 3 Da Nang, Vietnam 4 Hanoi, Vietnam 5 Lào Cai, Vietnam 108°21′E 105°51′E 103°57′E China 6 Haikou, Hainan 7 Chongzuo, Guangxi 8 Nanning, Guangxi 9 Guigang, Guangxi 10 Guangzhou, Guangdong 11 Shantou, Guangdong 12 Kunming, Yunnan 13 Guilin, Guangxi 14 Ganzhou, Jiangxi Altitude [m] Unangemeldet Heruntergeladen am | 10.04.17 15:37 J. Kovar-Eder / Acta Palaeobotanica 56(2): 329–440, 2016 MAT [°C] Longitude Climate station 440 Appendix 3. Climate data from the distribution region of the evergreen sclerophyllous broad-leaved forest sensu Wang (1961) as well as Platanus kerrii, Sloanea, and Ceratozamia today.

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Acta Palaeobotanica Early Oligocene plant diversity along the Upper Rhine Graben The fossil flora of Rauenberg Germany

Acta Palaeobotanica Early Oligocene plant diversity along the Upper Rhine Graben The fossil flora of Rauenberg Germany

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