New compression leaf material of Ceratozamia (Zamiaceae) has been recognised in the European Cenozoic. A leaflet of Ceratozamia floersheimensis (Engelhardt) Kvaček was recovered among unidentified material from the Oligocene of Trbovlje, former Trifail, Slovenia, housed in old collections of the Austrian Geological Survey, Vienna. It is similar in morphology and epidermal anatomy to other specimens previously studied from the lower Oligocene of Flörsheim, Germany and Budapest, Hungary. A fragmentary leaflet assigned to C. hofmannii Ettingsh. was recovered in the uppermost part of the Most Formation (Most Basin in North Bohemia, Czech Republic) and dated by magnetostratigraphy and cyclostratigraphy to CHRON C5Cn.3n, that is, the latest early Miocene. It yielded excellently preserved epidermal structures, permitting confirmation of the generic affinity and a more precise comparison with this lower Miocene species previously known from Austria (Münzenberg, Leoben Basin) and re-investigated earlier. Both the Oligocene and Miocene populations of Ceratozamia are based on isolated disarticulated leaflets matching some living representatives in the size and slender form of the leaflets. Such ceratozamias thrive today in extratropical areas near the present limits of distribution of the genus along the Sierra Madre Orientale in north-eastern Mexico, in particular C. microstrobila Vovides & J.D. Rees and others of the C. latifolia complex, as well as C. hildae G.P. Landry & M.C. Wilson (“bamboo cycad”). The occurrence of Ceratozamia suggests subtropical to warm-temperate, almost frostless climate and a high amount of precipitation. The accompanied fossil vegetation of both species corresponds well with the temperature regime. While the Oligocene species in Hungary probably thrived under sub-humid conditions, the remaining occurrences of fossil Ceratozamia were connected with humid evergreen to mixed-mesophytic forests.
Valid publication of new names of fossil plant taxa published since 1 January 1996 requires a diagnosis or description in English, besides other requirements included in the International Code of Nomenclature for algae, fungi, and plants (Melbourne Code) adopted by the Eighteenth International Botanical Congress, Melbourne, Australia, July 2011 (McNeill et al. 2012). In order to validate names published from the late Eocene flora of the Staré Sedlo Formation, North Bohemia, diagnosed only in German (Knobloch et al. 1996), English translations are provided, including references to the type material and further relevant information.
Zlatko Kvaček, Vasilis Teodoridis and Miroslav Radoň
The Oligocene palaeontological locality on Matrý Hill near Sebuzín in the České středohoří Mts., North Bohemia, belongs to the Děčín Formation and is dated to 30.8-24.7 Ma according to the regional stratigraphy. It has yielded, in addition to insects from the Heteroptera group, a fossil bee Apis petrefacta and palaeobatrachid frogs, also numerous plant remains. Their recovery began in 1996. The plant fossil assemblage consists mostly of leaf impressions, occasionally accompanied by casts or impressions of fruits. Noteworthy are records of a fern Woodwardia muensteriana, conifers Pinus cf. rigios, P. cf. hepios, Calocedrus suleticensis, Tetraclinis salicornioides, Torreya bilinica, cf. Cephalotaxus parvifolia and numerous angiosperms, e.g. Liriodendron haueri, Daphnogene cinnamomifolia, Platanus neptuni, Cercidiphyllum crenatum, Sloanea artocarpites, Ulmus pyramidalis, Celtis pirskenbergensis, Carya fragiliformis, C. quadrangula, Betula brongniartii, B. dryadum, Alnus rhenana, A. cf. kefersteinii, Carpinus grandis, Ostrya atlantidis, Acer crenatifolium, A. cf. palaeosaccharinum, A. integrilobum and Craigia bronnii. Several angiosperm foliage specimens of both monocots and dicots have not yet been identified to a particular genus and species. The fossil plant assemblage at Matrý corresponds to two vegetation types, i.e. a zonal riparian forest and zonal mixed mesophytic forest, as corroborated by the Integrated Plant Record vegetation analysis. The vegetation thrived under a humid climate, characterized by average values of MAT (13.4 °C), WMMT (23.8 °C), CMMT (3.6 °C) and MAP (1,117 mm). The Matry fossil flora is similar in composition to the flora of Žichov from the same Oligocene Děčín Formation in the České středohoří Mts.
New records of fruits of Banisteriaecarpum giganteum (Göppert) Kräusel from the middle and late Miocene of Austria and Hungary and the Pliocene of Romania regularly associated with foliage of Byttneriophyllum tiliifolium (A. Braun) Knobloch & Kvaček confirm previous views of Czeczott, Ţicleanu and others that the two organs belong to a single plant related to Malvaceae s.l. and not to Mapighiaceae, as previously assumed by Schenk, Kräusel and Kirchheimer. According to the fruit morphology it is closely similar to Tarrietia Blume (tropical SE Asia and Africa, sometimes included together with Argyrodendron F. Muel. to Heritiera Dryand. in Ait.), with which it partly shares habitats (swamp to riparian forests) and decidedly differs in foliage (leaves strongly asymmetric ovate vs symmetric simple ovate to elongate or palmately compound) and climatic requirements (warm temperate vs tropical conditions). Its pollen has not yet been firmly discriminated. The fossils so far assigned to Tarrietia from Europe must be excluded from this genus: Tarrietia hungarica Rásky from the early Oligocene of Hungary was assumed by Andreánszky as legume fruits (Machaerites hungaricus (Rásky) Andreánszky), Tarrietia germanica Rüffle from the early Miocene of Germany, according to fine venation pattern, may also represent a monospermic legume pod
The leaf remains described herein came from the oldest sites of the Cainozoic deposits in the Lower Rhine Embayment, located in the Siebengebirge Volcanic Field at the south-eastern border of this basin, in the area of Siebengebirge and vicinity. These revisited floras are bound to pre-volcanic siliciclastic facies of the Siebengebirge Mts., interpreted as marginal facies of the Köln Formation. Chronostratigraphically they are assigned to the late Oligocene (Chattian). The described leaf remains are partially compressions with preserved epidermal anatomy, and therefore highly useful for systematic determination of leaf impressions recovered from other localities of siliciclastic facies. On account of the epidermal characteristics of leaf compressions varying in gross morphology, the previously determined taxa Quercus goepperti, Laurus phoeboides, and Persea speciosa all fall into the abundantly represented Eotrigonobalanus furcinervis. The siliciclastic deposits originated in coastal and flood plain areas within fluviatile environments of variable deposition energy. Remains of Taxodium dubium, Eotrigonobalanus furcinervis, Populus germanica, and Daphnogene cinnamomifolia dominate among the recovered fossils. The general aspects of this plant assemblage correspond, together with their sedimentary settings, to riparian forest vegetation with mesophytic elements.
Accumulations of Osmunda (Plenasium) lignitum (Giebel) Stur fern foliage have been recently recovered in the Oligocene deposits of northern Italy at Cassinelle and western Germany at Niederpleis-1 (Sankt Augustin; Winterscheid 2006). They morphologically match other European records distributed from the Eocene to the Oligocene (– early Miocene). Material from both sites represents sterile pinnae compressions with preserved leaf anatomy. The data obtained on leaf cuticles correspond to most of the previous findings from Germany (Geiseltal, Seifhennersdorf, Haselbach) but cast doubts on some others based on dispersed cuticles (e.g. Zülpich). Comparison with living species of Osmunda (Plenasium) from east and south-east Asia revealed only minor differences in the epidermal structure and stress the close relationship with Osmunda banksiifolia (C. Presl) Kuhn distributed along valley streams from China, Japan to Indonesia, New Guinea and Philippines.
Zlatko Kvaček, Vasilis Teodoridis, Marianna Kováčová, Ján Schlögl and Viliam Sitár
A new plant assemblage of Cerová-Lieskové from Lower Miocene (Karpatian) deposits in the Vienna Basin (western Slovakia) is preserved in a relatively deep, upper-slope marine environment. Depositional conditions with high sedimentation rates allowed exceptional preservation of plant remains. The plant assemblage consists of (1) conifers represented by foliage of Pinus hepios and Tetraclinis salicornioides, a seed cone of Pinus cf. ornata, and by pollen of the Cupressaceae, Pinaceae, Pinus sp. and Cathaya sp., and (2) angiosperms represented by Cinnamomum polymorphum, Platanus neptuni, Potamogeton sp. and lauroid foliage, by pollen of Liquidambar sp., Engelhardia sp. and Craigia sp., and in particular by infructescences (so far interpreted as belonging to cereal ears). We validate genus and species assignments of the infructescences: they belong to Palaeotriticum Sitár, including P. mockii Sitár and P. carpaticum Sitár, and probably represent herbaceous monocots that inhabited coastal marshes, similar to the living grass Spartina. Similar infructescences occur in the Lower and Middle Miocene deposits of the Carpathian Foredeep (Slup in Moravia), Tunjice Hills (Žale in Slovenia), and probably also in the Swiss Molasse (Lausanne). This plant assemblage demonstrates that the paleovegetation was represented by evergreen woodland with pines and grasses in undergrowth, similar to vegetation inhabiting coastal brackish marshes today. It also indicates subtropical climatic conditions in the Vienna Basin (central Paratethys), similar to those implied by other coeval plant assemblages from Central Europe
Thomas Denk, Tuncay H. Güner, Zlatko Kvaček and Johannes M. Bouchal
The early Burdigalian (MN3) plant assemblage of the Güvem area (northwestern Central Anatolia) is preserved in lacustrine sediments of the Dereköy pyroclastics. Its age is well constrained by radiometric dates of basaltic rocks bracketing the pyroclastics, making the Güvem flora one of the extremely few precisely dated early Miocene floras in the Mediterranean region. The rich assemblage of impression fossils comprises ferns and fern allies (2 species), gymnosperms (12 spp.) and angiosperms (129 spp.). Ilex miodipyrena sp. nov. is described as a new fossil-species. The most diverse families in the assemblage are the Fagaceae with 12 taxa and the Fabaceae with 12 leaf morphotypes and one fruit taxon. Aquatic plants are represented by seven taxa, riparian (including palms) and swamp forest elements by >35 taxa, and lianas by three taxa (Smilax spp., Chaneya). The relatively large number of aquatic and riparian/swamp elements is congruent with the rich fish, amphibian and reptile record of the Güvem area. Another characteristic feature of the plant assemblage is the presence of various lobed leaves which show similarities with modern species of different families (e.g. Alangium, various Malvales). Trees and shrubs growing on well-drained soils and forming closed-canopy and open-canopy forests are the most diversified group (>70 taxa). In terms of number of specimens in the collection and based on field observations, by far the most abundant leaf fossils belong to evergreen oaks of Quercus drymeja and Q. mediterranea and to various types of foliage that cannot be assigned to a particular extant or extinct genus of Fagaceae. These sclerophyllous trees must have covered vast areas surrounding the wetlands that developed during the early Miocene in the Güvem Basin. Based on a recent reassessment of the ecology and taxonomic affinity of these trees, they are considered to reflect humid temperate climatic conditions but with a brief drier season during the winter months. These forests are more similar to the laurel forests of the southeastern United States and those stretching in a narrow belt south of the Himalayas to eastern central China. The large number of Fabaceae may indicate the presence of warm subtropical environments but this is difficult to assess, as they are known for having wide ecological ranges today and in the past. All in all, a larger part of the plant taxa point to forested vegetation. This is in agreement with previous palynological studies which detected only small amounts of herbaceous and grass pollen. Open patches of vegetation may have been restricted to river banks and to rocky areas in a volcanic landscape. The biogeographic patterns detected for the early Miocene of the Güvem assemblage are manifold; most taxa are widespread Northern Hemispheric elements. A substantial part of the species migrated from Asia into Europe during the (late) Paleogene and reached Anatolia during the early Miocene (Fagus, Paliurus, Chaneya, Ailanthus, Quercus kubinyii, Davallia haidingeri, Acer angustilobum, A. palaeosaccharinum). Fewer taxa may have been in Anatolia before they migrated to Europe (e.g. Nerium, Smilax miohavanensis, Quercus sosnowskyi). Finally, very few taxa are Anatolian endemics (e.g. Ilex miodipyrena).