Revision of Dendraena pinnatilobata Němejc from the Pennsylvanian of the Czech Republic


Authors: Frojdova J, Pšenička J, Bek J, Martinek K

Published in: Bulletin of Geosciences, volume 92, issue 1; pages: 75 - 94; Received 12 August 2016; Accepted in revised form 7 March 2017; Online 31 March 2017

Keywords: Dendraena, Anachoropteris, fern, Pennsylvanian, in situ spores,

full text (PDF, 4.31 MB)

Export to RIS



Dendraena pinnatilobata Němejc is a Pennsylvanian true fern, which is known from the Stradonice locality (Czech Republic) only. All specimens of Dendraena pinnatilobata are preserved as three-dimensionally in volcaniclastic of Whestone Horizon (Radnice Member, Kladno Formation). Revision of this species includes detail study of pinnae and pinnules morphology, sporangia, in situ spores and a rachial anatomy. Combination of several methods was used including camera lucida, maceration of sporangial/spores, SEM observation and cross sections of rachides for precise description of this species. D. pinnatilobata pinnae were borne on the rachides of the Anachoropteris robusta-type anatomy. This real organic connection between pinnae/pinnules (including reproductive organs) and rachial anatomy is rare in fossil record. This knowledge is very important for understanding of a leptosporangiate fern evolution. Sporangia of Dendraena pinnatilobata are annulate with band-lateral-upper type of annulus and in situ spores of the Microreticulatisporites harrisonii type, described for the first time, are unique for D. pinnatilobata. Based on this type of sporangia the species belongs to the leptosporangiate ferns. According to the petrological and sedimentological study of host sediments it is possible to infer that specimens of Dendraena were transported a short distance. Transport was very fast and fossils were deposited in supercritical conditions, which resulted in rapid burial of the plants and their excellent preservation. The plant host rock - Whetstone Horizon - also contains abundant volcaniclastic components. D. pinnatilobata grew in close proximity to the riverside or the peatland margins.


BARTHEL, M. 2016. The Lower Permian (Rotliegend) flora of the Dohlen Formation, Geologica Saxonica 61(2), 105-238.

BERTRAND, P. 1909. Études sur la frond Zygoptéridées. 306 pp. Ph.D. thesis. University of Lille, France.

BROUMISCHE, C. 1983. Les Fougeres sphénoptéridiennes du Bassin Houiller Sarro-Lorrain (Systematic-Stratigraphie). Publication de la Société Géologique du Nord 10, 1-480.

BROUMISCHE, C. 1986. Précisions sur les spores produites par quelques fougeres sphénoptéridiennes appartennant aux genres Boweria Kidston, Crossotheca Zeiller, Discopteris Stur, Myriotheca Zeiller et Urnatopteris Kidston. Revue de Paléobiologie 5(2), 231-248.

BROUMISCHE, C., COQUEL, R., LOBOZIAK, J.P., LOBOZIAK, S. & ŠETLÍK, J. 1985. Radiitheca dobranyana n. gen., n. sp., fougere pécoptéridienne du Carbonifere, productrice de spores de type Microreticulatisporites (Knox) Potonié et Kremp, 145-154.In DUTRO, T. JR. & PFEFFERKORN, H.W. (eds) Neuvieme Congres International de Stratigraphie et de Géologie du Carbonifere, Compte Rendu 5.

CORDA, A.J. 1845. Flora Protogaea, Beiträge zur Flora der Vorwelt. 128 pp. S. Calvary & Co., Berlin.

DETTMANN, M.E. 1963. Upper Mesozoic microfloras from south-eastern Australia. Proceedings of the Royal Society of Victoria 77, 1-148.

DIMICHELE, W.A. & PHILLIPS, T.L. 2002. The ecology of Paleozoic ferns. Review of Palaeobotany and Palynology 119, 143-159.View article

GALTIER, J. & PHILLIPS, T.L. 1996. Structure and evolutionary significance of Palaeozoic ferns, 417-433. In CAMUS, J.M., GIBBY, M. & JOHNS, R.J. (eds) Pteridology in Perspective. Royal Botanic Gardens, Kew.

GALTIER, J. & PHILLIPS, T.L. 2014. Evolutionary and ecological perspectives of Late Paleozoic ferns. Part III. Anachoropterid ferns (including Anachoropteris, Tubicaulis, the Sermayaceae, Kaplanopteridaceae and Psalixochlaenaceae). Review of Palaeobotany and Palynology 205, 31-73.View article

GOEPPERT, H.R. 1836. Die fossilen Farrnkrauter (System filicum fossilum). Verhandlungen der Kaiserlichen Leopoldisch-Carolinischen Akademie der Naturforscher 17(suppl.), 1-486.

HE, X.Z., WANG, S.J. & WANG, J. 2016. Chansitheca wudaensis (Gleicheniaceae, fern) from the early Permian Wuda Tuff Flora, Inner Mongolia. Palaeoworld 25(2), 199-211.View article

FROJDOVÁ, J. 2013. Leptosporangiate ferns from the Carboniferous basins of Bohemia, selected taxa. 61 pp. Master thesis, Faculty of Science, Charles University, Prague, Czech Republic.

FROJDOVÁ, J., PŠENIČKA, J., BEK, J. & CLEAL, J.C. 2017. Revision of the Pennsylvanian fern Boweria Kidston and the establishment of the new genus Kidstoniopteris, Review of Palaeobotany and Palynology 236, 33-58.View article

KIDSTON, R. 1911. Les végétaux houillers recueillis dans le Hainaut Belge et se trouvant dans les collections du Musée Royal d’Histoire Naturelle a Bruxelles. Mémoires du Musée royal d’histoire naturelle de Belgique 4(3), Mémoire no. 15, 1-282.

KIDSTON, R. 1923. Fossil plants of the Carboniferous rocks of Great Britain. Memoirs of the Geological Survey of Great Britain 2, 275-376.

KNOX, E.M. 1950. The spores of Lycopodium, Phylloglossum, Selaginella and Isoetes, and their value in the study of microfossils of Paleozoic age. Transactions of the Botanical Society of Edinburgh 35, 211-357.

LEHNERT, M.&WEIGAND, A. 2013. A proposal to distinguish several taxa in the Brazilian tree fern Cyathea corcovadensis (Cyatheaceae). Phytotaxa 155(1), 35-49.View article

NĚMEJC, F. 1930. Floristické poznámky ku stratigrafii některých uhelných pánviček mezi Rokycany a Berounem. Věštník státního geologického ústavu České republiky 6(1), 1-4.

NĚMEJC, F. 1934. On two new sphenopteris fructifications closely allied to Kidston’s Boweria (resp. Scott’s Pteridotheca) with remarks to some of the already known Palaeozoic fern sporangia. Věstník Královské České Společnosti Nauk, Tř. II, 1934, 1-18.

NĚMEJC, F. 1953a. Úvod do floristické stratigrafie kamenouhelných oblastí ČSR. 173 pp. Nakladatelství Československé akademie věd, Praha.

NĚMEJC, F. 1953b. Taxonomical studies on the fructifications of the Calamitaceae collected in the coal districts of Central Bohemia. Acta Musei Nationalis Pragae, series B, Historia Naturalis 9(1), 1-62.

NĚMEJC, F. 1963. Paleobotanika II. 523 pp. Československá akademie věd, Praha.

OPLUŠTIL, S. 2003. Sedimentation and Paleogeography of the Radnice Member (Duckmantian/Bolsovian) in the Kladno part of the Kladno-Rakovník Basin. Sborník Západočeského Muzea Plzeň, Příroda 102, 1-83.

OPLUŠTIL, S. 2005. Evolution of the Middle Westphalian river valley drainage system in central Bohemia (Czech Republic) and its palaeogeographic implication. Palaeogeography, Palaeoclimatology, Palaeoecology 222(3-4), 223-258.View article

OPLUŠTIL, S., PŠENIČKA, J., BEK, J., WANG, J., FENG, Z., LIBERTÍN, M., ŠIMŮNEK, Z., BUREŠ, J., DRÁBKOVÁ, J. 2014. T0 peat-forming plant assemblage preserved in growth position by volcanic ash-fall: A case study from the Middle Pennsylvanian of the Czech Republic. Bulletin of Geosciences 89(4), 773-818.View article

OPLUŠTIL, S., SCHMITZ, M., CLEAL, J.C. & MARTÍNEK, K. 2016. A review of the Middle-Late Pennsylvanian west European regional substages and floral biozones, and their correlation tot he Geological Time Scale based on new U-Pb ages. Earth-Science Reviews 154, 301-335.View article

PEŠEK, J. 2001. Středočeské a západočeské svrchnopaleozoické pánve, 10-73. In PEŠEK, J., HOLUB, V., JAROŠ, J., MALÝ, L., MARTÍNEK, K., PROUZA, V., SPUDIL, J. & TÁSLER, R. 2001. Geologie a ložiska svrchnopaleozoických limnických pánví České republiky. Český geologický ústav, Praha.

PEŠEK, J., HOLUB, V., JAROŠ, J., MALÝ, L.,MARTÍNEK, K., PROUZA, V., SPUDIL, J. & TÁSLER, R. 2001. Geologie a ložiska svrchnopaleozoických limnických pánví České republiky. 243 pp. Český geologický ústav, Praha.

PEŠEK, J. & MARTÍNEK, K. 2012. Observations concerning the thickness of rocks eroded between the Cambrian and Bolsovian (= Westphalian C) in Central and Western Bohemia. Folia Musei rerum naturalium Bohemiae occidentalis, Geologica et Paleobiologica 46(1-2), 1-14.

PETTIJOHN, F.J., POTTER, P.E. & SIEVER, R. 1973. Sand and Sandstone. 618 pp. Springer Verlag, Berlin.View article

PFEFFERKORN, H.W. 1978. Revision der Sphenopteriden Discopteris karwinensis Stur, Discopteris vuellersii Stur und der Gattung Discopteris Stur. Argumenta Palaeobotanica 5, 167-193.

POTONIÉ, R. & KREMP, G. 1954. Die Gattungen der Palaozoischen Sporae dispersae und ihre Stratigraphie. Geologisches Jahrbuch 69, 111-193.

POTONIÉ, R. & KREMP, G. 1955. Die Sporae dispersae des Ruhrkarbons ihre Morphographie und Stratigraphie mit Ausblicken auf Arten anderer Gebiete und Zeitabschnitte. Teil I. Palaeontographica 98B, 1-136.

PŠENIČKA, J. 2005. Taxonomy of Pennsylvanian-Permian ferns from coal Basins in the Czech Republic and Canada. 185 pp., 98 pls. Ph.D. thesis, Charles University, Prague, Czech Republic.

PŠENIČKA, J. & BEK, J. 2004. Tenchovia bulgarensis gen. et sp. nov., from the Pennsylvanian of the Dobrudzha Coalfield, Bulgaria. Geologica Balcanica 34(1-2), 69-76.

PUNT, W., HOEN, P.P., BLACKMORE, S., NILSSON, S. & THOMAS, A.L. 2007. Glossary of pollen and spore terminology. Review of Palaeobotany and Palynolology 143, 1-81.View article

REGE, R. 1920. Note su alcuni vegetali del Carbonifero della Cina. Atti della Societa italiana di scienze naturali e del Museo civico di storia naturale in Milano 59(3-4), 183-196.

REMY, R. & REMY,W. 1957. Durch Mazeration fertiler Farne des Palaozoikums gewonnene Sporen. Paläontologie Zeitschrift 31, 55-65.View article

RENAULT, B. 1896. Bassin Houiller Permien d’Autun et d’Épinac, Fascicule. IV, Flore Fossile, Deuxieme Partie, Texte ( Études des Gîtes Mineraux de la France). 578 pp. Imprimerie Nationale, Paris.

ROTHWELL, G.W. 1987. Complex Paleozoic Filicales in the evolutionary radiation of ferns. American Journal of Botany 74, 458-461.View article

SCHULZE, F. 1855. Uber das Vorkommen von wohlerhaltener Zellulose in Braunkohle undSteinkohle. Ber. Kgl. Preuss. Akad. Wiss. 676-678.

SCOTT, D.H. 1909. The Present Position of Palaezoic Botany. Progressus Rei Botanicae 1, 139-217.

SCOTT, D.H. 1920. Studies in fossil botany. Vol. I: Pteridophyta (3rd edition). 217-239 pp. A. & C. Black, London.View article

ŠIFTANCOVÁ, E. 2005. Rekonstrukce sedimentárního prostředí a provenience klastického materiálu radnických vrstev stradonického reliktu. Master thesis, Charles University, Prague, Czech Republic.

SMITH, A.H.V. & BUTTERWORTH, M.A. 1967. Miospores in the coal seams of the Carboniferous of Great Britain. Special Papers in Palaeontology 1, 1-324.

STÁRKOVÁ, M., ŠIMŮNEK, Z. & DRÁBKOVÁ, J. 2016. Geology of Carboniferous outlier between Lísek and Hýskov in paleobotanical and palynological context (Westphalian, Duckmantian). Geoscience Research Reports 49, 215-224.

STUR, D. 1883. Zur Morphologie und Systematik der Culm- und Carbonfarne. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Wien, Mathematisch-naturwissenschaftliche Classe 88 (I. Abt.), 633-846.

STUR, D. 1885. Beitrage zur Kenntniss der Flora der Vorwelt. II. Die Carbon-Flora der Schatzlarer Schichten. Abhandlundgen der Kaiserlich-königlichen geologischen Reichsanstalt 11, 1-418.

TAYLOR, T.N., TAYLOR, E.L. & KRINGS, M. 2009. Paleobotany: The Biology and Evolution of Fossil Plants (2nd edition). 1253 pp. Academic Press, Amsterdam.

TRIPLEHORN, D.M., STANTON, R.W., RUPPERT, L.F. & CROWLEY, S.S. 1991. Volcanic ash dispersed in the Wyodak-Anderson coal bed, Powder River Basin, Wyoming. Organic Geochemistry 17, 567-575.View article

ZEILLER, R. 1883. Fructifications de Fougeres du Terrain Houiller. Annales Scientifiques de la Nature, 6e série, Botanique 16, 177-209.

ZEILLER, R. 1897. Observation sur quelques Fougeres des dépôts houillers d’Asie Mineure. Bulletin de la Société de Botanique de France 44, 195-218.View article

ZEILLER, R. 1899. Étude sur le flore fossile du Bassin houiller d’Heraclee. Mémores de la Société Géologique de la France, Paléontologie 21, 1-91.