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Suttner, T.J., Kido, E., Königshof, P., Waters, J.A., Davis, L. & Messner, F. (Eds)

Planet Earth – In Deep Time Palaeozoic Series Devonian & Carboniferous

Suttner, T.J., Kido, E., Königshof, P., Waters, J.A., Davis, L. & Messner, F. (Eds)

With 201 coloured figures

Schweizerbart Science Publishers ∙ Stuttgart ∙ 2016 E Planet Earth – In Deep Time. Palaeozoic Series. Devonian & Carboniferous Suttner, T.J., Kido, E., Königshof, P., Waters, J.A., Davis, L. & Messner, F. (Eds)

We would be pleased to receive your comments on the content of this book: [email protected] Editors addresses: Suttner, Thomas J. – University of Graz, NAWI Graz, Institute for Earth Sciences (Geology and Palaeontology), Heinrichstrasse 26, 8010 Graz, Austria Email: [email protected] Kido, Erika – University of Graz, NAWI Graz, Institute for Earth Sciences (Geology and Palaeontology), Heinrichstrasse 26, 8010 Graz, Austria Email: [email protected] Königshof, Peter – Senckenberg, Forschungsinstitut und Naturmuseum Frankfurt, Senckenberganlage 25, 60325 Frankfurt, Germany Email: [email protected] Waters, Johnny A. – Department of Geology, Appalachian State University, Boone, NC 28608, USA Email: [email protected] Davis, Laura – 86 Tuscarora Ave, Beaufort, SC 29907, USA Email: [email protected] Messner, Fritz – Auenbruggergasse 8, 8073 Feldkirchen, Austria Email: [email protected]

Dust Cover: Artistic reconstruction of terrestrial ecosystem development from Silurian to Carboniferous in a transition-free time-series from left (back cover) to right (front cover). Computer generated collage by Messner F. (2007). Note for fossil collectors: Some fossil localities provided here are found in protected areas such as Geoparks, natural heritage or active military training ground, where access is restricted. Formal application and approval of access must absolutely be obtained before entry.

ISBN ebook (pdf) 978-3-510-65492-5 ISBN 978-3-510-65335-5 Information on this title: www.schweizerbart.com/9783510653355

© 2016 E. Schweizerbart‘sche Verlagsbuchhandlung (Nägele u. Obermiller), Stuttgart, Germany All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form means, electronic, mechanical photo copying, recording, or otherwise, without the prior written permissions of E. Schweizerbart‘sche Verlagsbuchhandlung, Stuttgart. Publisher: E. Schweizerbart‘sche Verlagsbuchhandlung (Nägele u. Obermiller) Johannesstraße 3A, 70176 Stuttgart, Germany [email protected] www.schweizerbart.de Printed on FSC-certified paper Layout: Thomas J. Suttner and Erika Kido Printed in Germany by Tutte Druckerei GmbH, Salzweg Preface

The International Geoscience Programme (IGCP) is different geological disciplines. There is no doubt that IGCP is one of UNESCO’s a cooperative enterprise of IUGS (International most successful scientific programmes. Union of Geosciences) and UNESCO (United Nations Educational, Scientific and Cultural Current and future climate change is one of society’s greatest challenges, in Organisation). IGCP is an international and multi- particular in terms of economy. It is well known that climate change affects disciplinary programme. It covers different fields in ecosystems, water supply and the frequency and magnitude of hazardous the Earth sciences and complements other phenomena among others. Furthermore, climate change is a global issue that can UNESCO and IUGS scientific programmes. IGCP be addressed only through international cooperation. Climate modelling is a began officially in 1972 when it was accepted at useful tool for a better understanding of climatological processes but cannot be the International Geological Congress at Montreal. done without the knowledge of organisms and changing palaeo-biodiversity. It is Attempts to create a global research effort began widely recognised that a temperature rise of about 1°C has occurred within the long before and there had been intensive work last 100 years and another rise is likely even if the amount is still in discussion. particularly by geologists in Europe (Austria, This shows how insecure is our knowledge of the complexity and functions of the Germany and UK) and Australia, to promote what was to become the most Earth’s climate system. The geologic and palaeontologic record of climate successful of IUGS/UNESCO-supported programmes. The primary aims of IGCP change is the best resource of information on the Earth’s climate system. It are to facilitate international collaboration among scientists in research on provides a record of trends in mean climate and changing palaeo-biodiversity. If geological problems, particularly between those individuals from more we can understand climate conditions that have existed during the history of industrialised and those from developing countries. The first Scientific Board recorded climate we gain a more fundamental understanding of the climate under the chairmanship of Sir Kingley Dunham (UK) with Simon van der Heide system than is attainable through studies based solely on modern conditions. acting for IUGS initiated some 28 projects. In the “early days” the scientific focus Furthermore, understanding climate change over the full variety of temporal and was primarily on basic science. The priority was the importance of accuracy and spatial scales recorded in the geologic sequences contributes to the knowledge definition of nomenclature to correlate whether geological data was stratigraphic of the fundamentals of climate dynamics. This knowledge is essential if we are to or otherwise. Identification and understanding of geological processes, model future climate with confidence. discovery and assessment of economic minerals, and the evolution of life are themes that are still reflected today, although the removal of the word In this context the IGCP 596 on “Climate Change and Biodiversity Patterns in the “correlation” shows a shift in emphasis away from the traditional basic geologic Mid-Palaeozoic” is an important contribution to a better understanding of climate projects. More recently, the emphasis changed towards societal-oriented themes. change in Earth history. The project is in line with UNESCO’s philosophy as is IGCP’s scientific objectives include: shown by the large number of contributions from different countries and languages. It is a nice example in terms of public outreach and provides  Improving our understanding of the geoscientific factors affecting the important information on both geological sections/areas and geosciences in global environment in order to improve human living conditions; general. The book will enhance visibility within the scientific community and  Developing more effective methods to find and sustainably exploit beyond the Earth Science community. natural resources of minerals, energy and groundwater;  Increasing the understanding of geological processes and concepts of global importance, including an emphasis on socially relevant issues; and  Improving standards, methods and techniques of carrying out geological research, including the transfer of geological geotechnological knowledge between industrialised and developing countries.

The objectives of IGCP are met through individual projects. Over the past Prof. Dr. Roland Oberhänsli decades, thousands of scientists have actively taken part in IGCP projects and for many of them the projects have been a gateway to a successful career. More IUGS President than 500 projects have resulted in thousands of scientific papers dealing with Contents

SeussB. and Mapes R.H. -The Finis Shalenear Jacksboro in Introduction 09 north-central Texas (Pennsylvanian)44 Kido E. and Suttner T.J. -Stratigraphy, Bioevents and Biodiversity10 Kido E. and Suttner T.J. -Continents and Plate Tectonics 12 Arabic translation of the IGCP 596 project summary 47 Königshof P. -What are scientificcollections and why they are important?14 (Ben Rahuma M.M. and Proust J.-N.) Planet Earth – In Deep Time: Devonian & Carboniferous 17 Libya Systematic overview of fossil groups Ben Rahuma M.M. and Proust J.-N.-The Awaynat WaninGroup in Trinajstic K. -Placoderms 20 Western Libya (Devonian)48 Kido E. -Tabulata and Rugosa22Morocco Suttner T.J. -Conodonts24El Hassani A. and Königshof P. -The Sabkhat Lafayrina Reef Complex Meyer-Berthaud B. -Cladoxylopsida 26 in the Tindouf Basin(Middle to Upper Devonian)50 Decombeix A.-L. -Lignophytes28El Hassani A. -The Ain Jemaa Reef Complex in the OulmèsArea, Stephenson C.A. -Isoetalean lycopsids30Moroccan Meseta (Middle to UpperDevonian)52 Kershaw S. -Stromatoporoids 32 SeussB. and Mapes R.H. -Cephalopoda 34 Bosnian translation of the IGCP 596 project summary 55 Kido E. and Brett C.E. -Arthropods 36 (Ćorić S.) Day J. -Rhynchonellata 38 Bosnia and Herzegovina Waters J.A. -Echinodermata 40 ĆorićS.and Hrvatović H. -The section of Mount Vranica in the Kido E. -Foraminifera and Radiolaria42Central Dinarides (Devonian)56 SeussB. and Nützel A. -Gastropoda 44 Bulgarian translation of the IGCP 596 project summary 59 English translation of the IGCP 596 project summary 19 (Boncheva I. and Sachanski V.) (Waters J.A.) Bulgaria Australia Boncheva I. and Sachanski V. -The section near Gigintsi Village in the Trinajstic K. -The Laidlaw Range in the Canning Basin(Upper Devonian)20Lyubash Monocline (Upper Devonian to Mississippian)60 George A. -Windjana Gorge in the Canning Basin(Upper Devonian)22 Trinajstic K., Hocking R. and Playton T.E. -The section at Burmese translation of the IGCP 596 project summary 63 Casey Falls, Canning Basin(Upper Devonian)24(Aung A.K.,BeckerR.T. and Myint K.K.) Meyer-Berthaud B. -The outcrop near Barraba in north-eastern Myanmar New South Wales (Upper Devonian)26Aung A.K. and KönigshofP. -The Pwepon Cave section near Decombeix A.-L. -The fossil locality near Dotswood in Kyadwinye Villageof theShan Plateau (MiddleDevonian)64 north-eastern Queensland (Mississippian)28 Ireland Chinese translation of the IGCP 596 project summary 67 Stephenson C.A. -Sandeel Bay on Hook Peninsula, (Wang Y.) County Wexford (Upper Devonian)30China United Kingdom Waters J.A., Chen X.Q., Suttner T.J. and Kido E. -The Boulongour Reservoir Kershaw S. - Hope’s Nose at Torbay in SouthwestEngland(Middle Devonian)32 section in northwestern Xinjiang (Upper Devonian)68 United States Wang Y. -The Naqing section in the Dian-Qian-Gui Basin, Guizhou Province Brett C.E. -The Seneca Stone Quarry in central New York (Pennsylvanian)70 (LowertoMiddle Devonian) 34 Brett C.E. -The cliffs of Lake Erie in western New York Czech translation of the IGCP 596 projectsummary 73 (Middle to Upper Devonian)36(Vodrážková S., Chadimová L., Tonarová P. and Vodrážka R.) Day J. -The Rockford Quarryin north-central Iowa Czech Republic (Upper Devonian)38Slavík L. -The Pragian GSSP at Velká Chuchlein the Waters J.A. -The outcrop at LakeCumberland in southern Kentuckey Prague Synform(Lower Devonian)74 (Mississippian)40Vodrážková S. -The Jirasek Quarry in the Prague Basin (Middle Devonian)76 Seuss B. -The Buckhorn AsphaltQuarry in southern Oklahoma Hladil J. and Poul I. -The Amphipora Limestone at Macocha Abyss (Pennsylvanian) 42 in the Moravian Karst (Middle Devonian)78 Dutch translation of the IGCP 596 project summary 81 Italian translation of the IGCP 596 projectsummary ------123 (Gouwy S.) (Corriga M.G. and Corradini C.) Belgium Italy Gouwy S. and Bultynck P. -The Couvinian stratotypeat Couvin Corriga M.G.,Corradini C. and Mossoni A. -The “Amphipora Limestone” at in the Ardennes (Middle Devonian)82Mount Zermula in the Carnic Alps (Middle Devonian) ------124 Simonetto L. and Corradini C. -The Riodel Museo section at French translation of the IGCP 596 project summary 85 Cason di Lanza, Carnic Alps (Pennsylvanian) ------126 (Nardin E.) Scanu G.G., Corriga M.G., Pillola G.L. and Corradini C. -The mining area Belgium near Iglesiasin southwestern Sardinia (Pennsylvanian) ------128 Da Silva A.C. and Boulvain F. -The La BoverieQuarry in the Ardennes (Upper Devonian)86Japanese translation of the IGCP 596 project summary ------131 Casier J.-G. -The Beauchâteau Quarry in the Ardennes (Kido E., Kurihara T. and Tsukada K.) (Upper Devonian)88Japan France Kurihara T. -The section in the Hikoroichiarea of the South Kitakami Belt Gouwy S. -The Charlemont Fortresssection at Givet in the Ardennes (LowertoMiddle Devonian) ------132 (Middle Devonian)90Tsukada K. and Manchuk N. -The section in the Fukuji-Hongo-Furukawa area Hubert B.L.M., Mistiaen B. and Pinte E. -The Parisienne Quarryin of the Hida Gaien belt (Devonian to Carboniferous) ------134 Boulonnais (Upper Devonian)92Higa K. and Nagai K. -The Akiyoshi Limestone of the Akiyoshi Accretionary MistiaenB. and Hubert B.L.M. -The Parc QuarryatEtrœungt in Terrane (Carboniferous to Permian) ------136 Avesnois(Upper Devonian)94 Latvian translation of theIGCP 596 project summary ------139 Senegal (Lukševičs E. and Stinkulis Ģ.) Sarr R. and Ngom P.M. -The Diana Malari Site in thenorthern Bové basin (Devonian)96Latvia Lukševičs E. and Stinkulis Ģ. -The Liepa Clay Pitin northern Latvia German translation of the IGCP 596 project summary 99 (Middle Devonian) ------140 (Suttner T.J. and Königshof P.) Lukševičs E. and Stinkulis Ģ.- The Remīne Quarryin central Latvia (UpperDevonian) ------142 Austria Lukševičs E. and Stinkulis Ģ. -The Salaspils Quarryin central Latvia Suttner T.J. and Kido E. -The Kirchfidisch Quarry in southern Burgenland (Upper Devonian) ------144 (LowerDevonian) ------100 Lukševičs E. and Stinkulis Ģ. -The Pavāri Site at the Ciecere River, Suttner T.J. and Kido E. -The Valentin Valley in the Carnic Alps south-western Latvia (UpperDevonian) ------146 (LowerDevonian) ------102 Kido E. and Suttner T.J. -The section at Mount Plabutsch in the Lithuanian translation of the IGCP 596 project summary ------149 Graz Palaeozoic(Middle Devonian) ------104 (Lazauskiene J.) Hubmann B. and Messner F. -The section at the Weisse Wand in theGrazPalaeozoic(Middle Devonian) ------106 Lithuania Kido E. and Suttner T.J. -The section at Mount Krone in the Lazauskiene J., Baliukevičius A. -The Petrašiūnai Quarryat the Carnic Alps (Pennsylvanian) ------108 Pakruojis district (Devonian) ------150 Germany Malaysian translation of the IGCP 596 projectsummary ------153 De BaetsK., Klug C. and Poschmann M. -The Hunsrück slate (Meor H.A.H.) in the Rhenish Massif (Lower Devonian) ------110 Königshof P. -The sequence of theLahn-Dillarea in the Malaysia Rhenish Massif (Middle to Upper Devonian) ------112 Hunter A.W., BashardinA. and Meor H.A.H. -The Pulau Langgun section Linnemann U. -The Bohlen section at Obernitz in the in the north-western Terrain(UpperDevonian) ------154 Saxo-Thuringian Zone (Devonian to Carboniferous) ------114 Hartkopf-Fröder C. -Wetlands in the Ruhr district (Pennsylvanian) ------116 Mongolian translation of the IGCP 596 project summary ------157 (Sersmaa G.) Hindi translationofthe IGCP 596 project summary ------119 Mongolia (Bhargava O.N.) Sersmaa G.,Ariunchimeg Ya., Kido E.,SuttnerT.J.,WatersJ.A., India Atwood J.W. and Webster G.D. -The Samnuuruul Formation section Bhargava O.N. and Draganits E. -The sedimentary sequence of the Baruunhuurai Terrane (Upper Devonian) ------158 of theTethyan Himalaya (Devonian) ------120 Ariunchimeg Ya.-The section at Hoshoot of the KhovdTerrane (Pennsylvanian)------160 Persian translation of the IGCP 596 project summary ------163 Siccardi A., Uriz N.J., Cingolani C.A. and Morel E.M.-The section of the (Sardar Abadi M.) Sierra de la Ventana Range (Middle Devonian) ------206 Iran Amenábar C.R. -The Chinguillos Group east of Blanco River in the Bahrami A. and Corradini C. -The Ghale-Kalaghu section in the Western Precordillera (Middle Devonian) ------208 Shotori Range (Devonian to Carboniferous) ------164 Amenábar C.R. -The section in theRio Blanco Basin of the SardarAbadi M.,Da Silva A.C. and Mossadegh H. -The Shahmirzad section Western Precordillera (Upper Devonian to Mississippian) ------210 in the Alborz Mountains (Mississippian) ------166 Spain Valenzuela-Rios J.I. and Liao J.-C.-The Compte-I section in the Polish translation of the IGCP 596 projectsummary ------169 Central Pyrenees (Lower Devonian) ------212 (Racki G.) Liao J.-C. and Valenzuela-Rios J.I. -The Renanué section in the Poland Central Pyrenees (Middle to UpperDevonian) ------214 Narkiewicz K. and Narkiewicz M. -The Zachełmie Quarry in the Uruguay western Holy Cross Mountains (MiddleDevonian) ------170 Uriz N.J., Siccardi A.,Portillo N., Cingolani C. andBlanco G. -The outcrop Racki G. -The Kowala Quarry near Kielce in the Holy Cross Mountains near the town of Blanquillo, Paraná Basin (Lower Devonian) ------216 (Upper Devonian) ------172 Wójcik K. -The Ostrówka Quarry in the Holy Cross Mountains (Upper Devonian) 174 Thai translation of the IGCP 596 projectsummary ------219 Skompski S. -The Ostrówka Quarry in the Holy Cross Mountains (Mississippian)176 (Charoentitirat T., Nantasin P. and Sardsud A.) Skompski S. -The Racławka Valley in the Cracow Region (Mississippian) ------178 Thailand Sardsud A.,Königshof P. and Charoentitirat T. -The Mae Sariang section Russian translation of the IGCP 596 project summary ------181 in northwestern Thailand (Upper Devonian) ------220 (Izokh N.G. and Obut O.T.) Russia Turkish translation of the IGCP 596 project summary ------223 ArtyushkovaO.V.and Mavrinskaya T.M. -The outcrop at the Belaya River (YalçınM.N.) in the Southern Urals (Lower Devonian) ------182 Turkey Artyushkova O.V. and Kulagina E.I. -The Irendyk Range in the Yalçın M.N. -The sequence of theArabian Plateand the Taurids (Devonian) ------224 Southern Urals (Upper Devonian) ------184 Yalçın M.N. -The sequence of the Pontids (Devonian) ------226 Artyushkova O.V. and Tagarieva R.Ch.-The Ryauzyak section in the Southern Urals (Upper Devonian) ------186 Ukrainian translation of the IGCP 596 projectsummary ------229 IzokhN.G.and Obut O.T. -The outcrop near Razdol’noe Village (Grytsenko V.) in the Rudny Altai (Upper Devonian) ------188 Ukraine Kulagina E.I.and Pazukhin V.N. -The Sikaza section in the GrytsenkoV.-The limestone quarry near Dzvenigorod Village Southern Urals (Mississippian) ------190 in the Dniester River area (Upper Silurian to Lower Devonian) ------230 Kulagina E.I. and Nikolaeva S.V. -The Kugarchi section in the GrytsenkoV.-The outcrop near Khoudykivtsy Village in the ZilairZone of the Southern Urals (Mississippian) ------192 Dniester River area (Lower Devonian) ------232 Kulagina E.I. and Nikolaeva S.V. -The Bashkirian stratotype at GrytsenkoV.-The outcrop near Nyrkiv Village in the Dniester area Yuryuzan River in the Southern Urals (Pennsylvanian) ------194 (LowerDevonian) ------234 Slovenian translation of the IGCP 596 project summary ------197 Vietnamese translation of the IGCP 596 project summary ------237 (Novak M.) (Ta HoaP.) Slovenia Vietnam Novak M. - The section at Dovžan Gorge in the Southern Doan Nhat T. and Ta HoaP.-The Lung Cu -Ma Le section of the Karavanke Mountains (Pennsylvanian) ------198 Dong Van Karst Plateau (Lower Devonian) ------238 Kolar-Jurkovšek T. and Jurkovšek B. -The outcrop at Doan Nhat T., Ta Hoa P. and Königshof P. -The section on Ljubljana Castle Hill (Pennsylvanian) ------200 Cát Bà Island (Mississippian) ------240 Spanish translation of the IGCP 596 project summary ------203 (Valenzuela-Rios J.I. and Liao J.-C.) Appendix ------243 Recommended Literature ------244 Argentina Figure captions ------252 CingolaniC.A., Uriz N.J., Manassero M.J. and Morel E.M. -The section near Authors ------258 Uspallata-Caracoles de Villavicencio,Precordillera(Lower Devonian) ------204 Acknowledgements ------261

Stratigraphy, Bioevents and Biodiversity

Sedimentary layers follow the basic law that the oldest layer is on catastrophic loss of taxonomic groups on Earth, occurred during bottom and youngest on top when the sequence of sediments is the end-Ordovician, Late Devonian, end-Permian, end-Triassic not overturned. Stratigraphy observes the inherent nature of and end-Cretaceous. The crises during the end-Ordovician and sediments, such as lithology, composition of fossils, geochemistry end-Frasnian (Late Devonian) probably resulted from intense (e.g., isotopic compositon), geophysics (e.g., magnetic rema- glaciation. Forty-nine percent and 35% of marine genera nence) and age. During much of Earth history, sedimentation has decreased respectively. The largest mass extinction since the been controlled by eustatic change of the sea level. The sea-level Cambrian is documented at the end-Permian leading to the loss of rise and fall during the depositional process results in certain 58% percent of marine genera. The effects of volcanism are cyclicity and is reflected in the sediments. The causes of sea-level considered as one of the causes of this event. The Central fluctuations are the change of seawater volume, which is Atlantic Magmatic Province (CAMP), one of the major volcanic associated with the development of glaciers for example, and the provinces developed during the Triassic, erupted voluminous long term aspect—the change of alignment of continents and flood basalts and is considered a major cause responsible for the oceanic basins. end-Triassic crisis. During this crisis, 40% of all marine genera went extinct. The end-Cretaceous mass extinction was linked with The sea-level fluctuations, the division of Ecologic Evolutionary impacts and major volcanic eruptions. Thirty-nine percent of Units, the marine family diversity, and the time of well-known mass marine genera disappeared. Extinction events destroyed common extinction events (Big Five) are shown in the left figure of the next community organizations. Reorganization of communities was the page. In terms of diversity, the marine fossil record is divided into consequence. Many clades moved into new environments, which three Successive Evolutionary Faunas (Cambrian, Palaeozoic and resulted in new association of species or replacement by invading Modern), which have distinctive compositions and progressively species. increasing diversity. During the intervals of Evolutionary Faunas, new organism-groups brought new morphologic adaptations into The Devonian was a relatively warm period with an acme in the existing communities. During their radiation, communities diversity, size and latitudinal distribution of reefs during the increased in diversity and complexity through geological time. Middle Devonian. Nonetheless, apart from the end-Frasnian mass Nine intervals shown (two in Cambrian, four in the remaining part extinction, several climate perturbations that resulted in additional of the Palaeozoic, and three in Modern) conform to Ecologic more-or-less-severe biotic events have been documented (Figure Evolutionary Units (EEUs), which are assigned as the ecologic next page, right side). During the Carboniferous (except for late subdivision of Evolutionary Faunas. Visean and late Gzhelian) extensive pulses of glaciation coincided with high-frequency sea-level fluctuations observed in deposits Five large mass extinctions in the Phanerozoic, which caused globally.

Kido E. and Suttner T.J.

Continents and Plate Tectonics

The Earth’s crust moves in mm- to cm-scale per year. This Pangaea. The collision of continents or microcontinents, and the movement results in diastrophism (i.e., deformation of earth’s accretion of terranes that went with the formation of Pangaea surface) which is folding and faulting within the crust. The cause caused mountain-building activity that produced the Appalachian of diastrophism was earlier explained by Alfred Wegener in 1912, Mountains, Ural Mountains and the Rhenish Massif among other when he proposed the theory of continental drift (The Origin of mountain chains. Continents and Oceans; Wegener, 1915). It was advanced by the theory of mantle convection in 1928 and by the theory of seafloor In this book, palaeogeographic reconstructions by Ronald Blakey spreading in the early sixties. These theories were summarised (NAU Geology) are used. The location of the countries that appear and developed as plate tectonics by John Tuzu Wilson after the in the book is plotted on the modern map (right page) with the late 1960s. Plate tectonics is a theory that explains the colour code to each country provided below. The location of fossil diastrophism based on the motions of tectonic plates on the sites in those countries is plotted on Devonian and Carboniferous earth’s surface, which consist of continental and oceanic palaeogeographic maps in each contribution. lithosphere. Following the plate tectonics theory, configuration of ocean basins and continents through geological time are reconstructed based on palaeomagnetic data and palaeogeographic interpretation of fossil occurrences. The palaeogeographic maps have been improved and published by several scientists (e.g., Scotese and McKerrow, 1990; Golonka, 2002; Kiessling and others, 2003; Paleomap Project by Scotese: http://www.scotese.com; Reconstructing the Ancient Earth by Blakey: http://cpgeo- systems.com/paleomaps.html). During the Devonian, the supercontinent Gondwana covered vast areas of the southern hemisphere and the continent Siberia was located in the northern hemisphere. Other major continents such as Laurussia lay between them. Gondwana was divided from eastern Laurussia by the Proto-Tethys Ocean and from the microcontinents (e.g., Avalonia), which existed in the south of Laurussia by the Rheic Ocean. The Carboniferous was the time when formation of the supercontinent Pangaea began. Two major oceans were spread in this period: the Palaeo-Tethys Ocean (a successor of the Proto-Tethys Ocean) and the Panthalassa Ocean that surrounded

Kido E. and Suttner T.J.

What are scientific collections and why they are important?

IGCP 596 is specifically interested in the interaction between studies and for all areas that depend on systematics, such as climate change and biodiversity in the Devonian and Carboni- biostratigraphy, palaeobiogeography and reconstruction of the ferous Periods (419 – 299 million years ago) when the terrestrial history of life. ecosystems experienced a biodiversity boom and oceanic ecosystems suffered catastrophic extinctions. Furthermore, palaeontological collections provide important resources for palaeontological education at all levels (e.g., graduate Fossils are used to investigate these changes in biodiversity of school, education in and out of museums). Many institutions host different periods in Earth history. Palaeontology has become a key huge numbers of organisms, particularly museums such as the component particularly in palaeoclimatic research. Together with Smithonian (USA); Natural History Museum, London (U.K.); sedimentary rocks, fossils are the repository of nearly all American Museum of Natural History, New York (USA); and the palaeoclimatic data, especially quantitative data. For instance, Senckenberg Museum, Frankfurt (Germany). The unique stable isotope geochemical interpretations of palaeoclimate are importance is also based on the fact that smaller institutions and most commonly measured from fossils. Understanding the universities have decided to rid themselves of their collections. evolution, palaeobiology, and palaeoecology of organisms is Access to scientific collections is a prerequisite to investigate the crucial to palaeoclimate interpretations of stable isotope records. extinct fauna and flora. Palaeontological collections will benefit For example, vertical or horizontal migrations during ontogeny, as enormously from greatly increased accessibility to information well as seasonality in reproduction may have consequences for provided by computer technology. Therefore, related to this estimates of ocean temperatures from marine microfossils. The project, a network of taxonomic workers will help to update the palaeobiology and palaeoecology of organisms is also very systematics of Mid-Palaeozoic terrestrial and marine organisms. important to interpretations of shifts in palaeobiogeographic These datasets will be made available to the public by using patterns in terms of palaeoclimatic change. existing e-infrastructures such as the Paleobiology Database.

Therefore, natural history collections are vital scientific research Based on the scientific collections – a key component in palaeo- tools. Palaeontological collections are fundamental for all ntological research and research infrastructure – the project palaeontological information and they are essential for research in might help give understanding to our present-day situation and all areas of palaeontology, from taphonomy to palaeoecology to climate change in the future by documentation of Mid-Palaeozoic evolutionary palaeobiology. They are also essential for systematic climate change and its effect on biodiversity.

Königshof P.

International Geoscience Programme (IGCP) IGCP Project 596 (2011-2015) Climate change and biodiversity patterns in the Mid-Palaeozoic

The face of Planet Earth has changed significantly through ecosystems unknown before the Devonian Period. The success of geologic time. The configuration of continents and oceans and the terrestrial invaders, as documented by the fossil record, organisms that inhabited them are very different from those we culminated with the development of vast forests consisting of see today. Dynamic processes still active today, such as plate tree-like forms like Calamites (Order Equisetales), lycophyte trees tectonics and climate change, have shaped the earth’s surface (e.g. Lepidodendron, Sigillaria) and other rooted plants that and impacted biodiversity patterns from the beginning. IGCP 596 covered huge areas during the Carboniferous. That unique rise is specifically interested in the interaction between climate among land plants and the formation of top-soil led to distinctive change and biodiversity in the Devonian and Carboniferous changes in environmental conditions. Based on proxy-data, we Periods (419 – 299 million years ago) when the terrestrial can show that the rapid rise of land plants was coupled with ecosystems experienced a biodiversity boom and oceanic strongly decreasing atmospheric CO2 values from 4000 ppm to ecosystems suffered catastrophic extinctions. nearly present day values of about 350 ppm during the latest Devonian. Increased weathering activity and soil formation by Greenhouse climates dominated the Early and Middle Devonian rooted plants lead to intensified run-off and changed water (419 – 383 Ma) world, but changed to icehouse conditions in the chemistry, which seriously affected marine communities globally. Late Devonian (383 – 359 Ma). The early Carboniferous world was relatively warm until cooling in the early late Carboniferous (323 – The tectonic and climate history of the Devonian and 299 Ma) resulted in a huge polar ice shield in the southern Carboniferous as well as the novelty of soil-formation due to the hemisphere that covered most of Gondwana. The Mid-Palaeozoic explosion of life on land, and other processes, some of which are was also a time of very high plate tectonic activity that caused not yet fully understood, are linked with a series of ecological major palaeogeographic changes. During the Devonian two turnovers and extinction events primarily in the oceans. Results of supercontinents, Euramerica and Gondwana, together with this project should help to clarify whether climate change (e.g. Siberia formed the biggest landmasses of our planet. They interaction of CO2 and temperature) from greenhouse conditions successively amalgamated into the supercontinent Pangaea during the Early-Middle Devonian to icehouse conditions during during the late Carboniferous. As the continental landmass grew, the Late Devonian – early Carboniferous represents a major vascular plants, arthropods, hexapods and first tetrapods spread trigger for variations in biodiversity or if a combination of multiple on land. Their radiation formed the base of new terrestrial factors is responsible for such changes.

Waters J.A.

The Devonian Reef sediments of the Canning Basin in the northwest of Placoderms (Phylum Chordata) ruled the oceans for 70 Western Australia are divided into a number of different facies types million years from the Silurian until the end of the Devonian. representing different palaeoenvironments. The early Frasnian Gogo Formation is world renown for the exceptional preservation of the pelagic The fossil record of placoderms is parti- marine vertebrate and invertebrate fauna. The formation was cularly good as the head and thorax deposited in the deeper waters of a barrier reef along the were covered with plates northern margin of Gondwana. made of dermal bone. Their diverse body The Gogo Formation comprises fine black forms and different shales interbedded with thinly laminated arrangements of the carbonate nodules. The fossils form the skull plates distinguish nucleus of the nodules and are composed of different placoderm taxa. mostly pelagic organisms. They include The exceptionnal preservation invertebrates such as phyllocarid and conca- of the placoderms from the Gogo vicarid crustaceans, radiolarians, sponges, Formation have been important to conodonts and a single eurypterid. However, understanding the evolution of the novel the area is best known for the fish fauna, morphology in vertebrates. Placoderms repre- which is dominated by placoderms (extinct sent the first vertebrates to have jaws with armoured plated fish; Figure: Eastmanosteus teeth, paired pectoral and pelvic girdles, calliaspis), but also includes osteichthyans (boney fish), and internal fertilization with live rare sharks and acanthodians, marine lungfishes and birth. The first evidence of tetrapodamorphs. In contrast, the bottom fauna is restricted sexual dimorphism, where with only rare molluscs and brachiopods. The fauna is preserved males looked different to three dimensionally and includes original bone and preservation of soft females, was observed by tissues. The restricted benthic fauna suggests low oxygen levels at Watson in ptyctodonts sediment/water interface and the three dimensional preservation from Scotland, however it indicates a high sedimentation rate resulting in rapid burial. Over 50 fish was not until 2008 when a species were recorded from this site with the placoderms as the most single embryo attached by an umbilical cord was discovered from the abundant and diverse group. The preservation of a complete ecosystem Gogo Formation that internal fertilization with live birth was confirmed in provides important information about the past species diversity and the ptyctodont Materpiscsi attenbourough, name after Sir David allows a more accurate reconstruction of the extinct reef community. Attenbourough.

Trinajstic K. Trinajstic K.

One of the best known localities in the reefs of the Canning Basin is Tabulata and Rugosa (Phylum Cnidaria, Class Anthozoa, Windjana Gorge, which was eroded by the Lennard River. The limestone Subclass Zoantharia) were sessile marine organisms that forming the gorge walls record ~10 million years of reef growth (middle produced calcified skeletons. These coral groups appeared inde- Frasnian to early Famennian), with movement of the reef margin pendently in the Ordovician and became extinct at the end of the recording the response of the reef builders to relative Permian. Palaeozoic corals are found in different facies changes in sea level. including limestone, calcarerous siltstone and shales. In the shallow marine environment, tabulates, rugosans and The Canning Basin reefs grew continuously through an other calcified metazoans produced extensive carbonate important time of global change with a cooling platforms. In the Canning Basin, corals were important greenhouse Earth and a major set of global extinctions in components of the platform lagoons and, to a lesser extent, the late Devonian. The major Frasnian reef builders were the reef margins. stromatoporoids and calcimicrobes and they belonged to a diverse ecosystem with sponges, corals and skeletal The Tabulata developed exclusively compound forms. organisms such as crinoids, ammonoids, nautiloids, Massive coralla with common skeletal tissue, massive trilobites, brachiopods, molluscs and armoured fishes cerioid coralla with mural pore (small holes developed in (Figure: Frasnian reef fabric with platy stromatoporoids, the wall, which link neighbouring corallites) and branching calcimicrobes beneath and finely layered red sediment coralla were dominant. The Rugosa developed mainly with fossil fragments). Towards the end of Frasnian, the solitary corallites, but produced also colonial forms of Canning Basin reefs were affected by global environ- fasciculate, massive cerioid and amural growth-type. mental changes. Significantly, this region was characte- Tabulate corals were characterised by a simple skeleton rised by tectonic activity causing fault movements and with well-developed tabulae and inconspicuous septa. In rapid relative sea-level changes. Anoxic ocean water does contrast, rugose corals had septa typically consisting of a not appear to have been a factor in the faunal changes in major and minor one, with diversely developed tabulae, the Canning Basin at this time. Local environmental dissepiments and axial structures. changes and cooling global climate severely impacted the stromatoporoids and they were largely extinct before the end of Frasnian. Although several coral species were encrusting after their planulae The microbes were not so obviously affected if at all, and unlike many (larvae) had settled on hard grounds, the vast majority of corals lived on other regions around the world, they continued to build major reefs in the soft substrates when mature. Coral diversity was highest in warm, Famennian. Overall, global cooling and tectonic quiescence in the shallow environment. Compared with recent zooxanthellate scleractinian Canning Basin eventually led to the demise of reefs before the end of corals, Tabulata and Rugosa are thought not to have had symbiotic Famennian. algae.

George A. Kido E.

The Virgin Hills Formation (early Frasnian – middle Famennian) in the Conodonts belong to a group of extinct chordates that Canning Basin reef complexes is a characteristically reddish silty occurred from the Cambrian through the late Triassic. They limestone well known for its diverse and beautifully preserved marine were first described in a monograph on Silurian fish by Christian fossils. It was deposited in marginal slope and basin environments in Heinrich Pander (Riga, Lithuania) in 1856. Since then, the commonly front of a steep reefal margin in warm tropical seas. The formation is found millimetre-sized, coniform to complex-shaped tooth-like exposed along the SE Lennard Shelf. A section through Casey Falls elements of this fossil group has been assigned to several groups, records two significant events; the Frasnian/Famennian (F/F) extinction ranging from plants up to marine vertebrates. To which group and the end of reef building in the Canning Basin – both marked by conodonts finally belong is not clarified yet. Currently it seems most significant faunal overturns. The Virgin Hills Formation shows no reasonable to assign them to marine basal chordates. In 1983 the significant facies change across the F/F extinction boundary, although first “conodont-animal” was described from the Mississippian the water depth is interpreted to be in the hundreds of metres. The Granton Sandstone, Scotland. That specimen is unique, as it is a fossils include stromatolites, microbes and algae; invertebrates such complete body-fossil of 40.5 mm in length and less than 1.8 mm in as conodonts (Figure) crinoids, tentaculites, and brachiopods; and width, with soft tissue preserved all over. Conodont elements are rare vertebrates such as scales from jawless fish (thelodonts), arranged in situ in the head region of the specimen. Another shark teeth and scales and placoderm plates. body fossil from late Ordovician deposits of South Africa There are local occurrences of stromolitic boundstone reached a length of at least 400 mm. encrustation within the Virgin Hills Formation. The control of their timing and distribution is speculative, but imply relatively starved Based on soft tissue analysis, conodonts are characterised conditions in a distal setting, which the silty nodular carbonates that by having two large eyes, a posterior and caudal fin dominate the Virgin Hills Formation also reflect. There is a major supported by rays, v-shaped muscle segments and a palaeoenvironmental change with reef building ceasing in the complexly arranged feeding apparatus including 15 to 19 trachytera conodont Zone. Overlying the Virgin Hills Formation is the discrete elements. These elements consist of apatite (calcium late Famennian Bugal Gap Limestone. After an apparently brief sea level phosphate) and are usually found disarticulated, as fused fall the late Famennian – Tournaisian Fairfield Group was deposited on a clusters in coprolites of marine predators, or as natural marine carbonate ramp that draped the relic reefal profiles. Changes in assemblages with apatite elements in situ with soft tissue lithology and depositional environment were accompanied by a major preservation. It is proposed that they were active swimmers near the change in the fauna at this time. In the Fairfield Group conodonts are rare, sea-bottom. Whether they were predators or parasites, or had a but brachiopods, ostracods and corals are abundant. There is also an saprophage or grasping life style is uncertain. Although this group still increase in the type and variety of shark teeth and scales, and in the has some systematic and palaeobiological uncertainties, its apatite teeth and scales of boney fish (palaeoniscoids), however the placoderms elements are one of the most important tools used for microfossil become extinct at the Devonian – Carbonifeous (359 Ma) boundary. biostratigraphy during the entire Palaeozoic and early Mesozoic.

Trinajstic K., Hocking R. and Playton T.E. Suttner T.J.

The late Devonian floras of Australia are commonly called Cladoxylopsida represent an extinct group of plants “Leptophloeum floras” due to the overwhelming occurrence of affiliated to the ferns. It ranges from the Eifelian (Middle compression/impression remains of the arborescent Devonian) to the Mississippian (early Carboniferous) and is lycopsid genus Leptophloeum. A few localities distributed worldwide. It is known to have evolved yielding anatomically preserved fossils, however, the earliest trees, in the Middle Devonian. indicate that Australian floras were more diverse than suggested by this designation. Anatomically, the Cladoxylopsida The plant beds occurring close to the are the earliest plants to show locality of Barraba, in north-eastern a complex vascular system New South Wales, have yielded consisting of a network of several plants affiliated to the ferns, primary vascular strands among which is Polyxylon australe. and little, if any, secondary This endemic species of Clado- xylem, as in modern ferns. xylopsida is represented by anato- This pattern is generally very mically preserved stems measu- attractive when stems are ring up to 25 mm in diameter observed in transverse section. (Figure: transverse section and stem). Analysis of the branching pattern in The plant beds belong to the Mandowa Polyxylon australe Mudstone, which consists of thinly shows that this bedded and massive mudstone plant had widely alternating with thin beds of spaced bran- siltstone and fine grained ches borne on sandstone. The depositio- slender stems and nal environment was a resembled a liana rather distal marine shelf to conti- than a small tree. This suggests nental slope. The plant that, by the end of the Devonian, the beds in the Mandowa Mud- Cladoxylopsida had evolved a range of elaborate stone at Barraba are Famen- growth forms and contributed to the formation of nian (late Devonian) in age. complex plant ecosystems.

Meyer-Berthaud B. Meyer-Berthaud B.