Zlatko Kvaček

The Integrated Plant Record: An Essential Tool For Reconstructing Neogene Zonal Vegetation In Europe

Abstract A semi-quantitative evaluation method has been developed to map integrated fossil plant records (leaf, fruit, and pollen assemblages) in terms of zonal vegetation. It incorporates taxonomy, physiognomy, and autecological features of the fossil plants. To derive vegetation types, the final stage of this method involves a quantitative evaluation. The maps presented here are based on 198 organ assemblages from 173 localities and stratigraphic levels of the Miocene–early Pliocene across Europe. They cover three time intervals: (1) 4–7 Ma (Messinian–middle Zanclean; late Miocene–early Pliocene); (2) 8.5– 12 Ma (latest Serravallian–middle Tortonian); and (3) 14–17 Ma (late Burdigalian–late Langhian). We also look at the interval 13–14 Ma (late Langhian–early Serravallian). They not only show vegetation types but offer more detailed information on the composition of these associations (proportions of major components). The major zonal vegetation types recognized here are broad-leaved deciduous forests, mixed mesophytic forests, broad-leaved evergreen forests, subhumid sclerophyllous forests, open woodlands, and xeric grasslands. Between 14 Ma and 17 Ma, broad-leaved evergreen forests were widespread, and subhumid sclerophyllous forests were more strongly represented in western parts of Central Europe than in eastern parts. In the interval 8.5–12 Ma, broad-leaved deciduous forests largely replaced the evergreen forests, although these were still present in some refugia (e.g., the Lower Rhine embayment and northern Balkan peninsula), and first records of xeric grasslands are found along the northern margin of the Black Sea. The more discontinuous record available from 4 Ma to 7 Ma indicates that broad-leaved evergreen forests still occurred in northern parts of the Mediterranean (northern Italy, Balkan peninsula), while open woodlands increasingly appeared in central and southern parts of Italy and Greece.

Paleobotanical studies in Fagaceae of the European Tertiary

A systematic reassessment of megafossil records of Fagaceae in Central Europe has been undertaken on the basis of leaf cuticular characters. The oldest representatives date back to the Eocene: Quercus subhercynica spec. nova, Dryophyllum furcinerve (Rossm.) Schmalh., Trigonobalanopsis rhamnoides (Rossm.) gen. & comb. nov. In the Oligocene other members of extant genera appear: Quercus rhenana (Weyl. & Kilpp.) Knobloch & Kvacek, Fagus attenuata Goepp., Lithocarpus saxonicus spec. nova. In the Neogene these ancient taxa (except in Fagus lineage), are gradually replaced by deciduous species of Quercus and Castanea. Trigonobalanus and Castanopsis are recorded by fruits (or wood) only.

Bı́lina: a window on Early Miocene marshland environments

Abstract The Early Miocene deposits at Bilina (northern Bohemia) have yielded a wealth of plant fossils belonging to various sedimentary settings. The flora includes ferns (8 species), conifers (6 species) and angiosperms (more than 80 species). The modern Arcto-Tertiary element predominates. Vegetation of coal-forming swamps, flood plains/levees, shallow waters and crystalline uplands can be reconstructed. Aquatic plant horizons dominated by Salvinia contain unusual angiosperm herbs belonging to the Araceae, Lemnaceae, Hydrocharitaceae and ?Scrophulariaceae. Swamp forest was dominated by the Taxodiaceae, swampy broad-leaved trees ( Nyssa , Alnus , Myrica ), monocotyledons (palms, Zingiberaceae, Stratiotes ) and ferns. Riparian forest included Taxodium , hygrophilous deciduous broad-leaved woody dicotyledons ( Ulmus , Alnus , Fraxinus , Acer ), various lianas and a few mesophytic elements ( Parrotia , Zelkova , Podocarpium ). Crystalline uplands were covered by mixed forest of acidic soils with Pinus , Myrica , Comptonia , Trigonobalanopsis , Lauraceae. The palaeoclimate can be characterized as optimal warm-temperate, very humid, perhaps with a lower humidity during the winter.

Fossil Leaves of the Araceae from the European Eocene and Notes on Other Aroid Fossils

A comparative study of leaf morphology and venation patterns of extant Araceae has been undertaken in order to define groups matching aroid fossil leaves from the Eocene of Central Europe. As a result, four morphogenera have been recognized: Araceophyllum Kräusel emend. Wilde, Z. Kvaček & Bogner (subfamily Pothoideae); Araciphyllites Wilde, Z. Kvaček & Bogner, gen. nov. (subfamilies Monsteroideae and Aroideae); Caladiosoma Berry (tribes Colocasieae and Caladieae); and Nitophyllites Iljinskaja emend. Fedotov (tribes Peltandreae and Arophyteae). The leaf fossils studied from the Eocene sites at Messel and Eckfeld (Germany) and Kučlín (Czech Republic) have been included in four species: Araciphyllites tertiarius (Engelhardt) Wilde, Z. Kvaček & Bogner, comb. nov. (common at Messel, rare at Eckfeld); Araciphyllites schaarschmidtii Wilde, Z. Kvaček & Bogner, sp. nov. (a single specimen at Messel); Caladiosoma messelense Wilde, Z. Kvaček & Bogner, sp. nov. (rare at Messel); and Nitophyllites bohemicus Wilde, Z. Kvaček & Bogner, sp. nov. (a single fragment at Kučlín). Outside Europe, other Tertiary aroid foliage species that we have revised nomenclaturally include Araciphyllites tobleri (Kräusel) Wilde, Z. Kvaček & Bogner, comb. nov. (Neogene of Sumatra), and Nitophyllites limnestis (Dilcher & Daghlian) Wilde, Z. Kvaček & Bogner, comb. nov. (Eocene of the Claiborne Formation, Tennessee). Critical notes are also provided concerning other Cenozoic records of Araceae and their affinities, like Nitophyllites zaisanicus Iljinskaja emend. Fedotov (late Paleocene‐Eocene of Kazakhstan and the Russian Far East) and Peltandra primaeva Hickey (Eocene of the Golden Valley Formation, North Dakota). The mentioned four species of aroid foliage add considerably to the reliable fossil record of the family that now contains ca. 10 species based on leaves. Their nearest living relatives suggest a (para)tropical to subtropical climate that is well in accordance with the paleogeographic position and the accompanying floras of the respective localities. Except for the two species of Araciphyllites from Messel, they all obviously represent helophytes of aquatic/hemiaquatic or muddy/swampy sites. Except for Caladiosoma miocenicum, which is suggested to be of Neogene age, all of the fossil leaves of Araceae are mostly of Eocene age, suggesting a considerable diversity of Araceae as early as in the early Paleogene. We caution against taking for granted all fossil records whose names imply affinities with the Araceae. Many of them are excluded from the family or are at least doubtful. Others, mainly based on reproductive structures or whole plants, contribute important information for understanding of the fossil history of aroids.

Comparing Early to Middle Miocene floras and probable vegetation types of Oberdorf N Voitsberg (Austria), Bohemia (Czech Republic), and Wackersdorf (Germany).

The Early Miocene vegetation of western Styria, Austria, is reconstructed on the basis of detailed investigations of leaves and diaspores from the mining area Oberdorf N Voitsberg. In this paper, the flora and probable vegetation are compared with other assemblages of similar age from the Czech Republic and Germany to elucidate the diversity of wetland and mesophytic plant communities. The floristic composition and the vegetational units represented in Oberdorf are compared to the middle part of the Most Formation (M Most Fm), Cypris Shale, Mydlovary Formation (Mydlovary Fm) of Bohemia, and the Wackersdorf mining area of Bavaria. Among these sites, Oberdorf is extremely poor in hydrophytes and reed-like monocotyledons, indicating rare pond-like habitats as well as possibly more closed swamp forest conditions than, e.g. in the M Most Fm. In Oberdorf, relatively high pH-values in the lignite-forming swamps and the possible scarcity of acidic, nutrient-poor (sandy) soils have probably influenced the floristic composition of the azonal communities. The peat-forming associations in all regions share the abundance of Glyptostrobus europaeus. However, distinct differences in the accompanying elements, such as Taxodium dubium (present and abundant only in the M Most Fm) indicate even stronger floristic variability of Early(/Middle) Miocene peat-forming and riparian plant communities than previously expected. The virtual absence of Pinus, Engelhardia, Comptonia, and probably also of Quercus kubinyii/Quercus drymeja in the megafossil record of Oberdorf can possibly be explained by the edaphic conditions. The abundance of Sequoia abietina (absent in all the other compared sites), which we assign to riparian (and mesophytic) forests in Oberdorf, may have been favoured by rich alluvial soils.In the mesophytic associations, thermophilous elements of the Lauraceae, Mastixiaceae, Symplocaceae, and Rutaceae are diverse. Usually they include Trigonobalanopsis rhamnoides/Trigonobalanopsis exacantha, an evergreen Fagaceae. This assemblage type corresponds with the Younger Mastixioid Flora sensu Mai. The Younger Mastixioid Flora is best developed in Wackersdorf, less distinct in Oberdorf, and likely in the Cypris Shale and Mydlovary Fm. It is not traceable in the M Most Fm.

TRIFOLIOLATE LEAVES OF PLATANUS BELLA (HEER) COMB. N. FROM THE PALEOCENE OF NORTH AMERICA, GREENLAND, AND ASIA AND THEIR RELATIONSHIPS AMONG EXTINCT AND EXTANT PLATANACEAE

Trifoliolate leaves of Platanus bella (Heer) comb. n., a species previously known only from the Paleocene of western Greenland, are newly recognized from the Paleocene of northern Wyoming, U.S.A., and Altai of Xinjiang Province, northwestern China, indicating that the species was circumboreal in the early Tertiary. Epidermal anatomy preserved in specimens from all three areas confirms that these compound leaves belong to the Platanaceae. Platanus bella (Heer) comb. n. differs from modern species of Platanus but resembles the European Tertiary species Platanus fraxinifolia (Johnson & Gilmore) Walther and Platanus neptuni (Ettingshausen) Bůžek, Holý & Z. Kvaček, in the presence of large peltate glandular trichomes. We erect a new subgenus, Glandulosa, to accommodate P. bella, P. fraxinifolia, and P. neptuni. Each of these species possesses similar leaf epidermal structure, with the characteristic platanaceous stomatal apparatus and compound hair bases. In addition, the fossils bear peltate glandular trichomes on the epidermal surfaces that are not known among extant Platanus species. Reproductive structures linked to P. neptuni indicate that subgenus Glandulosa is properly placed in the Platanaceae but that it is a distinct clade from those of the extant subgenera Platanus and Castaneophyllum Leroy. We also review the status of the fossil genera Debeya and Dewalquea, to which some Cretaceous and Tertiary leaves of similar morphology have been placed, and reject the use of either of these names to accommodate leaves of subgenus Glandulosa.

Limnobiophyllum Krassilov — a fossil link between the Araceae and the Lemnaceae

Abstract Limnobiophyllum Krassilov includes fossil free-floating stoloniferous plants each with one or two sessile suborbicular to reniform leaves of different size as well as numerous simple and one or two longerbranched roots on a reduced main stem. The venation consists of campylodromous curved primaries, up to 14 in number, among which irregular reticulate veins of higher orders can be preserved. There are no signs of lateral pouches characteristic of the Lemnaceae. However, aerenchyma and pigment cells are well developed in Limnobiophyllum . Two species are recognized - Limnobiophyllum scutatum (Dawson) Krassilov (latest Cretaceous to Oligocene of western North America and Palaeocene of East Asia) and Limnobiophyllum expansum (Heer) Kvacek, comb. n. (Miocene of Europe). Turion-like bodies are associated with L. scutatum . No fruits have been found in connection, but numerous isolated ribbed seeds are associated with L. expansum . They resemble some Araceae and also Lemna L. and Spirodela Schleid. Such seeds are known as Lemnospermum Nikitin from the Tertiary of Eurasia. Limnobiophyllum resembles in some respects Spirodela , but it is larger and the roots and the habit are more like young plants of Pistia L. (Araceae). It differs from Hydromystria Mey. (i.e. Limnobium Rich.) by venation, root system and habit; the Lemnaceae, although similar, differ by lateral pouches of leafy fronds and the lack of branched roots and higher-order veins. Limnobiophyllum is considered as an extinct link related to Pistia (Araceae), from which Spirodela (Lemnaceae) may have evolved by reduction.

CONES, SEEDS, AND FOLIAGE OF TETRACLINIS SALICORNIOIDES (CUPRESSACEAE) FROM THE OLIGOCENE AND MIOCENE OF WESTERN NORTH AMERICA: A GEOGRAPHIC EXTENSION OF THE EUROPEAN TERTIARY SPECIES

The cupressaceous genus Tetraclinis is recognized from the Oligocene and Miocene of western North America on the basis of co‐occurring seed cones, seeds, and foliage branches. Morphological and anatomical comparisons with the two previously recognized European Tertiary species indicate that the North American specimens are morphologically inseparable from Tetraclinis salicornioides (Unger) Kvaček. The North American taxon is treated as a new variety, T. salicornioides (Unger) Kvaček var. praedecurrens (Knowlton) comb. et stat. nov., and is distinguished from the European representatives, T. salicornioides (Unger) Kvaček var. salicornioides, by slight anatomical differences in the leaf epidermis. Although cones and seeds of the fossil species are closely similar to those of extant Tetraclinis articulata, the foliage is more “spreading,” composed of flattened segments with fused facial and lateral leaves that are apparently adaptive for a more mesic climate. The recognition of T. salicornioides in western North America along with the absence of Tetraclinis in the fossil and recent flora of eastern Asia provide evidence for communication of the species across the North Atlantic during the early or middle Tertiary.

Fossil Fruits of Pteleaecarpum Weyland-Tiliaceous, not Sapindaceous

Tertiary fruit remains of Pteleaecarpum, which we recently interpreted as valves of winged capsules of an extinct sapindaceous genus, are now recognized as belonging to a living genus of Tiliaceae. The fossil fruits, which are common in the Tertiary of western North America, Europe, and Asia, are virtually identical in morphology and anatomy to fruits of Craigia W. W. Smith and Evans, an extant tiliaceous genus of southern China. Accordingly, we now present the following new combinations for fossil taxa: Craigia bronnii (Unger) and Craigia oregonensis (Arnold).

Twig with attached leaves, fruits and seeds of Decodon (Lythraceae) from the Lower Miocene of northern Bohemia, and implications for the identification of detached leaves and seeds

Abstract A twig fragment with attached verticillate leaves and capsules with intact seeds was recovered from the Lower Miocene lignitic clay in the Bilina strip mine (northern Bohemia). The seeds correspond to those known in dispersed condition as Decodon gibbosus (E.M. Reid) E.M. Reid in Nikitin, while the foliage matches that which has been called Ficus multinervis Heer, and sometimes Apocynophyllum helveticum Heer. While the determination of the seeds is safe, the identity of the isolated foliage may be less secure. Therefore the name Decodon gibbosus is given preference to designate the whole plant. This fossil species differs from the only extant species D. verticillatus (L.) Ell. of eastern North America by narrower triangular shape of seeds in transverse section, the thicker spongy layer of the testa on the dorsal side, and by long petiolate leaves that are more variable in shape (from narrow linear-lanceolate to broadly lanceolate) with more densely spaced secondaries and intersecondaries. The leaf cuticular structure matches that of the glabrous form of D. verticillatus. The associated plant assemblage suggests aquatic to swampy environment typical of the extant relative.