TRIBUTE TO CHARLES DARWIN AND
BERNISSART IGUANODONS:

New Perspectives on Vertebrate Evolution and Early
Cretaceous Ecosystems

BRUSSELS 2009

EDITORS: PASCAL GODEFROIT & OLIVIER LAMBERT

Darwin-Bernissart meeting, Brussels, February 9-13, 2009

FOREWORD

We are pleased to welcome you to the Extraordinary Meeting of the European Association of
Vertebrate Palaeontologists (EAVP) `A tribute to Charles Darwin and Bernissart Iguanodons: New
perspectives on Vertebrate Evolution and Early Cretaceous Ecosystems', hosted in Brussels by the
Royal Belgian Institute of Natural Sciences (RBINS). February 12, 2009 is a key-date for European
palaeontologists. Indeed, we are to celebrate the 200th birthday of Charles Darwin, the Father of the
Evolution Theory, and also the official opening of a new Gallery of Evolution in the capital of Europe.
Moreover, February 2009 corresponds to the closure of the `Bernissart Iguanodons' Year', marking
the 130th Anniversary of the discovery of these jewels of the RBINS. We are delighted that so many
palaeontologists accepted our invitation to gather in Brussels and to celebrate these unique events,
through highlighting recent developments in their specific research fields.

The first section of the meeting is devoted to Vertebrate Palaeontology and Evolution, with a first day
(Monday, February 9) composed of short talks dealing with various fields of vertebrate palaeontology.
The second day (Tuesday, February 10) is organized differently, composed of longer plenary lectures
tackling some of the main themes developed in the new Gallery of Evolution of the RBINS (Cambrian
radiations, the first vertebrates, Palaeozoic fish evolution, the first land plants, the early tetrapods,
Cenozoic birds, the evolutionary history of the cetaceans, domestication...). These talks should be
addressed to a wider audience, including university teachers and researchers in the fields of biology
and geology, graduate students, scientific journalists...

The second section deals with new perspectives on Bernissart and other Early Cretaceous terrestrial
ecosystems. The first day of this section (Wednesday, February 11) is held at Bernissart, the village
where the famous Iguanodons were discovered in 1878. It is mainly devoted to synthetic
communications about iguanodontids and Early Cretaceous ecosystems in Belgium, Europe, and the
world. The second day (Friday, February 13), held at the RBINS, is composed of more technical talks
about recent discoveries in Early Cretaceous terrestrial localities of the Bernissart area and Eurasia,
and about some aspects of dinosaur anatomy.

Two fieldtrips are organized at the occasion of this meeting. On Thursday February 12, Gaël Clément
(Muséum national d'Histoire naturelle, Paris) and Cyrille Prestianni (Ulg, Liège) lead a visit to the
Upper Devonian locality of Strud, in the Namur Province. The estuarine to fluviatile sediments of this
small quarry yielded a rich Famennian vertebrate fauna, among which placoderms, acanthodians,
actinopterygians, sarcopterygian fishes and early tetrapods (related to the famous Ichthyostega),
together with outstanding plant and arthropod material. The visit of a remarkable fossil collection
(fishes, echinoderms, brachiopods...) from the Lower Visean 'Marbre Noir de Denée', at the
Maredsous Abbey, is organized by Eric Groessens (Geological Survey of Belgium, RBINS). On
Saturday February 14, Johan Yans (FUNDP, Namur) leads an excursion at Hautrage, in the Hainaut
Province. This site is estimated to be contemporaneous with the Iguanodon Cran at Bernissart. The
drilling of a series of boreholes in this "Iguanodon Cran" has recently provided the impetus for a
renewed study of the Lower Cretaceous sites of Belgium, including Hautrage, which yielded a very
rich fossil plant assemblage. Not only leaves and fruits were found, but also big pieces of trunk, some
of them charcoalified, and all deposited in an alluvial context.

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
We would like to express our sincere thanks to the people and organisations, which have contributed
to this meeting. We are greatly indebted to the Belgian Science Policy, our main sponsor, and to its
President, Philippe Mettens, who encouraged the organisation of this symposium. The General
Director of the RBINS, Camille Pisani, provided support, advice and facilities. Willem De Vos,
Kareen Goldfeder and the Communication staff of the RBINS, as well as members of the Department
of Palaeontology, tirelessly helped us in various practical aspects of the organisation and with helpful
hints. Bertrand Panier (RBINS) designed the conference logo and the cover of this volume. The Mayor
Roger Vanderstraeten, Annette Cornelis, Corinne Detrain, the Tourism Office of Bernissart, the Local
Development Agency of Bernissart, and the Geological Circle of Hainaut efficiently provided meeting
support and facilities at Bernissart. Marie-Céline Buchy generously proposed two grants from the
Raymonde Rivoallan Fund, which allowed us to offer grants to several palaeontologists presenting
communications at the meeting. Eric Groessens and Paul Van Genabeek organized the visit at the
Maredsous Abbey.

THE ORGANIZING COMMITTEE,

Pascal Godefroit,
Olivier Lambert,
Etienne Steurbaut,
Johan Yans,
Gaël Clément,
Mona Court-Picon,
Adriano Vandersypen

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009

iv

Darwin-Bernissart meeting, Brussels, February 9-13, 2009

TRIBUTE TO CHARLES DARWIN AND
BERNISSART IGUANODONS:

New Perspectives on Vertebrate Evolution and Early
Cretaceous Ecosystems

BRUSSELS 2009

Programme, Abstracts and Field Trips Guidebook

Programme ................................................................................ 1
Abstracts .................................................................................... 7
Field Trips Guidebook ................................................................. 103

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
PROGRAMME

Sunday February 8, 2009 - Registration of the participants

15.00-18.00 Arrival of the conference participants and registration at the front desk of the Museum of the
Royal Belgian Institute of Natural Sciences (rue Vautier 29, 1000 Bruxelles)

Monday February 9, 2009 - Scientific presentations for the `Vertebrate Palaeontology and
Evolution' section, at the Royal Belgian Institute of Natural Sciences (large auditorium)

Session 1 - Moderator: A. Blieck

09.00-10.00 Arrival of the conference participants and registration at the front desk of the Museum of the
Royal Belgian Institute of Natural Sciences (rue Vautier 29, 1000 Bruxelles)

10.00-10.15 Welcome speech by Etienne Steurbaut, Head of the Section Fossil Vertebrates at the RBINS

10.15-10.35 Fack F. & Derycke C. Diet evidences on Viséan crushing-tooth (Cochliodus) from Soignies
(Belgium)
10.35-10.55 Olive, S., Goujet, D. & Lelièvre, H. A new saoudian taxon of Acanthothoraci (Placodermi):
questions on plates homologies and growth
10.55-11.15 Cuny, G., Cavin, L. & Suteethorn, V. Specialized dentition in Lower Cretaceous freshwater
hybodont sharks

11.15-11.35 Coffee Break

Session 2 - Moderator: J. Clack

11.35-11.55 Sorin Baciu, D., Miclu , C., Grdianu, I. & Niculi, M. Fish fauna from Oligocene bituminous
marls of Marginal Folds Nappe, East Carpathians, Romania
11.55-12.15 Marjanovi, D. & Laurin, M. A reevaluation of the evidence supporting an unorthodox hypothesis
on the origin of the extant amphibians
12.15-12.35 Fischer, V., Guiomar, M. & Godefroit, P. New data on the palaeobiogeography of Toarcian
(Lower Jurassic) ichthyosaurs
12.35-12.55 Elgin, R. A., Prondvai, E., Frey, E. & Krüger, W. Pterosaur flight dynamics and the evolution of
flight style within the lineage: initial experimental findings

12.55-14.15 Lunch

Session 3 - Moderator: J.-M. Mazin

14.15-14.35 Rabi, M. Presence of Sebecosuchian crocodyliforms in the Late Cretaceous of Europe
14.35-14.55 si, A. The Late Cretaceous continental vertebrate fauna from Iharkút, western Hungary a
review of the newest results
14.55-15.15 Gilissen, E. & Smith, T. Early mammalian brain diversity: insights from multituberculates
15.15-15.35 Hautier, L. & Saksiri, S. Description of the masticatory apparatus of Laonastes aenigmamus
(Rodentia, Diatomyidae), new insights into the evolution of hystricognathy
15.35-15.55 Serdyuk, N.V. & Vasiliev, S.K. Mammalian fauna of Hyena den cave (Altai, Russia)

15.55-16.15 Coffee Break

Session 4 - Moderator: G. Clément

16.15-16.35 Hoch, E. Contribution to the evolutionary history of the beaked whales
16.35-16.55 Lambert, O., Bianucci, G. , de Muizon, C. & Dooley, A. C. Jr. New stem sperm whales from the
Miocene of Peru
16.55-17.15 Steeman, M. E. The problematic fossil baleen whales from Belgium
17.15-18.00 Plenary lecture by Simon Conway Morris. Darwin's compass: why evolution is predictable

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
19.00
Welcome Drink and Beer Party in a typical old-style bar, La Bécasse, in the centre of Brussels
(Rue de Tabora 11, 1000 Bruxelles)

Tuesday February 10, 2009 - Plenary lectures: 'Key Moments of Evolution in a Nutshell', at the
Royal Belgian Institute of Natural Sciences (large auditorium)

09.00-09.15 Welcome speech by Camille Pisani, General Director of the RBINS

Session 1 - Moderator: A. Milner

09.15-10.00 Mayr, G. Composition and biogeographic affinities of the middle Eocene Messel avifauna
10.00-10.45 Bultynck, P. To be or not to be: can conodonts be vertebrates?

10.45-11.05 Coffee Break

11.05-11.50 Blieck, A. Palaeozoic biodiversity, ecosystems and evolution: the case of Ordovician to Devonian
vertebrate-dominated assemblages
11.50-12.35 Gerrienne, P. Early land plant evolution: a tremendous success story
12.35-13.20 Clément, G. & Prestianni, C. Biodiversity of the Devonian tetrapod-bearing locality of Strud,
Belgium

13.20-14.30 Lunch

Session 2 - Moderator: E. Hoch

14.30-15.15 Clack, J. What's new in the world of Devonian tetrapods?
15.15-16.00 Milner, A. R. Carboniferous amphibians and reptiles experimentation and diversification
16.00-16.45 Bardet, N. Les faunes de reptiles marins du Crétacé terminal du Maroc

16.45-17.05 Coffee Break

17.05-17.50 de Muizon, C & Geisler, J. H. From land to sea, the early evolution of cetaceans
17.50-18.35 Germonpré, M. The domestication of the dog during the Upper Palaeolithic
18.35-19.20 Vignaud, P. Origine de l'Homme: les nouvelles du Tchad

20.00
Symposium Dinner at the Royal Belgian Institute of Natural Sciences

Wednesday February 11, 2009 - Scientific presentations for the `Bernissart and other Early
Cretaceous Terrestrial Ecosystems' section, at the 'salle communale' in Harchies

09.00-10.00 Bus travel to Bernissart-Harchies

10.00-10.10 Welcome speech by Philippe Mettens, President of the Belgian Federal Science Policy Office

Session 1 - Moderator: P. Taquet

10.10-10.30 Buffetaut, E. L'importance historique de la découverte des Iguanodons de Bernissart
10.30-10.50 Yans, J., Andreu, B., Baele, J.-M., Cornet, C., de Ricqlès, A., Dejax, J., Dupuis, C., Galbrun, B.,
Gerards, T., Gerrienne, P., Godefroit, P., Gomez, B., Gosselin, R., Leduc, T., Petit, G., Pons, D.,
Preat, A., Robaszinski, F., Schnyder, J., Smith, T., Spagna, P., Steurbaut, E., Taverne, L.,
Tshibangu, J.-P., Van Itterbeeck, J., Vandycke, S. & Vanneste, C. Integrated results of the new
material from the 2002-2003 boreholes in Bernissart (Mons Basin, Belgium)
10.50-11.10 de Ricqlès, A., Godefroit, P. & Yans, J. Vertebrate remains in the 2003 Bernissart drill:
histological assessment

11.10-11.30 Coffee Break

Session 2 - Moderator: D. Norman

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
11.30-11.50 Pouech, J. & Mazin, J.-M. Faunal assemblages at the Jurassic/Cretaceous boundary: comparison
between some West European continental sites
11.50-12.10 Buffetaut, E. Early Cretaceous dinosaurs and pterosaurs from the eastern Paris Basin
12.10-12.30 Wilde, V. & Riegel, W. The terrestrial Lower Cretaceous of Germany ("Wealden")

12.30-14.30 Lunch and visit of the discovery place of the Bernissart Iguanodons

Session 3 - Moderator: E. Buffetaut

14.30-14.50 Pereda Superbiola, X., Ruiz-Omeñaca, J. I., Canudo, J. I., Torcida, F., Sanz, J. L. Lower
Cretaceous dinosaurs of Spain: an overview based on skeletal remains
14.50-15.10 Fuentes Vidarte, C., Meijide Calvo, M., Meijide Fuentes, F. & Meijide Fuentes, M. A new
Iguanodon species (Dinosauria, Ornithischia) from the Lower Cretaceous of Soria (Spain)
15.10-15.30 Posmosanu, E. Early Cretaceous ornithopod dinosaurs from Romania
15.30-15.50 Taquet, P. The African cousins of the Iguanodons

15.50-16.10 Coffee Break

Session 4 - Moderator: P. Godefroit

16.10-16.30 Pan, R., Wang, Y. & Wang, X. A new iguanodontian dinosaur from the Early Cretaceous Yixian
Formation of Western Liaoning, China
16.30-16.50 Roolf, C. The attempted theft of dinosaur skeletons during the German occupation of Belgium
1914-1918 as a topic of the history of science and the history of total war
16.50-17.10 Goussard, F. Zalmoxes, Rhabdodon and Rhabdodontidae: How many genera and how many
species?
17.10-17-30 Mazin, J.-M., Pouech, J. & Raslan-Loubatié, J. The Bernissartids: common European crushing
crocodiles
17.30-18.15 Plenary lecture by David B. Norman. Iguanodontians from the Wealden of Britain and
Europe

18.15-19.30 Speech by Mr. Roger Vanderstraeten, Mayor of Bernissart

Visit of the `Musée de l'Iguanodon' at Bernissart by Corinne Detrain

Special auction organized to raise funds for the Raymonde Rivoallan Fund

19.30 -22.00 Dinner at Bernissart-Harchies

22.00-23.00 Bus travel to Brussels

Thursday February 12, 2009 - Field trip in Strud, poster session and official inauguration of the
new Gallery of Evolution at the Museum of Natural Sciences

09.00-10.15 Bus travel to Strud

10.15-11.00 Visit to the Upper Devonian (Famennian) quarry of Strud, led by Gaël Clément (MNHN, Paris)
and Cyrille Prestianni (ULg, Liège)

12.30-13-30 Lunch at the Abbey of Maredsous

13.30-14.30 Visit to the collection of fossils from the Carboniferous (Early Visean) 'Marbre Noir de Denée', at
the Abbey, led by Eric Groessens (RBINS, Brussels)

14.30-15.45 Bus travel back to Brussels

15.45-16.05 Coffee break

16.05-17.30 Poster session

17.30
Official inauguration of the new Gallery of Evolution and dinner at the RBINS

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
Friday February 13, 2009 - Scientific presentations for the `Bernissart and other Early
Cretaceous Terrestrial Ecosystems' section at the Royal Belgian Institute of Natural Sciences
(large auditorium)

Session 1 - Moderator: P. Bultynck

09.00-09.20 Christian, A., Dzemski, G. & Möller, J.-T. Neck utilization and feeding strategy in sauropod
dinosaurs
09.20-09.40 Vandycke, S. & Spagna, P. Crustal tectonic control of the Early Cretaceous deposits in the Mons
Basin (Belgium)
09.40-10.00 Licour, L. Mississipian formations and Cretaceous tectonics : The impact of the actual geothermal
reservoir genesis on the past landscapes of the Mons Basin
10.00-10.20 Quinif, Y. The karstic phenomenon of Bernissart pit and the geomorphologic situation in the
Mesozoic times

10.20-10.40 Coffee Break

Session 2 - Moderator: J. Le Loeuff

10.40-11.00 Martin, T., Yans, J., Dupuis, C., Spagna, P. & Kaufmann, O. 3D modelling of the paleozoic top
surface in the Bernissart area and integration of data from boreholes drilled in the "Cran aux
iguanodons"
11.00-11.20 Masure, E. & Yans, J. Stratigraphy of the Haine Groupe ("Meule" sediments) overlying the
Wealden facies in the BER 3 Borehole (Bernissart, Mons Basin, Belgium)
11.20-11.40 Schnyder, J., Dejax, J., Keppens, E., Nguyen Tu, T. T., Spagna, P., Riboulleau, A. & Yans, J.
Organic matter characterization and organic carbon isotopes record in the Early Cretaceous
lacustrine setting of Bernissart (Belgium)
11.40-12.00 Yans, J., Dejax, J., Gerards, T., Gerrienne, P., Spagna, P. & Keppens, E. Carbon isotopes on
woody material from Wealden facies of Hautrage (Mons Basin, Belgium)
12.00-12.20 Gomez, B., Yans, J., Gillot, T., Spagna, P., Coiffard, C. & Daviero-Gomez, V. From past
collections of Bernissart to recent collections of Hautrage: new taxonomical and environmental
insights

12.20-14.00 Lunch

Session 3 - Moderator: J. Yans

14.00-14.20 Baele, J.-M., Quesnel, F., Bourdillon, C. & Dupuis, C. Cretaceous coastal environments in the
Mons Basin: Evidences from Cenomanian deposits in the Bettrechies Quarry
14.20-14.40 Dejax, J., Pons, D. & Yans J. Palynological overview of Wealden facies sediments from Belgium
and Northern France
14.40-15.00 Leduc, T. diagenesis of the fossil bones of Iguanodon bernissartensis
15.00-15.20 Lauters, P, Coudyzer, W. Vercauteren, M. & Godefroit, P. Iguanodon's brain and perspectives on
ornithopod evolution
15.20-15.40 de Santisteban, C. & Santos-Cubedo, A. The record of climate and eustatic changes during the
lower cretaceous in the "Arcillas de Morella Formation" (Spain)

15.40-16.00 Coffee Break

Session 4 - Moderator: E. Steurbaut

16.00-16.20 Moreno-Azanza, M., Gasca, J. M. & Canudo, J. I. New data on the Valanginian-Hauterivian
reptile ootaxa of the Iberian Range (NE of Spain)
16.20-16.40 Le Loeuff, J. Insularity and dwarfism in Late Cretaceous European dinosaurs
16.40-17.00 Stein, K. & Sander, M. Quantifying growth rates in island dwarf sauropods
17.00-17.20 Mallison, H. Rearing for food? Kinetic/dynamic modelling of bipedal/tripodal poses in sauropod
dinosaurs
17.20-17.40 Godefroit, P., Jin, L. & Chen, J. A new middle Cretaceous vertebrate locality from Jilin Province,
northeastern China
17.40-17.50 Closing of the meeting

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
Saturday February 14, 2009 - Field trip in Hautrage

Important: Pair of boots and a windcheater are strongly recommended because of the humid environment of the
visited site.

09.00-10.00
Bus travel from Brussels to Hautrage

10.00-12.00
Visit of the Hautrage Quarry, led by Johan Yans (FUNDP, Namur)

12.00-13.00
Bus travel from Hautrage to Brussels

List of posters

The posters will be on display in a special room during the whole meeting time. Therefore, the posters will be
available during talks, coffee breaks and lunch. A special poster session is scheduled for Thursday 12, between
16.05 and 17.30, just before the official inauguration of the new Gallery of Evolution, when the authors are
asked to be present besides their poster.

Badiola, A., Canudo, J. I. & Cuenca-Bescós, G. New Early Cretaceous multituberculate fossils from the Iberian
Peninsula
Bianucci, G., Gatt, M. & Catanzariti, R. Oligo-Miocene odontocetes from the Maltese islands: stratigraphical
distribution and evolution
Chanthasit, P. The Braincase of Rhabdodon (Dinosauria: Ornithopoda): new specimens from the late Cretaceous
of southern France
Codrea, V., Godefroit, P., Smith, T. & Jipa-Murzea, C. Maastrichtian land vertebrates in Rusca Montan Basin
(Romania).
Dumont, M., Borbely, A., Kostka, A. & Pyzalla, A. Transmission electron microscopy of sauropod bones: a look
inside the nanostructure
Falconnet, J. Permocarboniferous amniotes from France: a state of the art and perspective
Garcia, G., Godefroit, P., Smith, T., Van Itterbeeck, J., Valentin, X. & Codrea, V. Amniotic eggshells from the
Haeg Basin (Upper Cretaceous, Romania)
Garcia, G., Andossa, L., Mackaye, H.-T., Vignaud, P. & Brunet, M. Chelonians from the Late Miocene of Chad
(the hominid site TM 266)
Gasca, J. M., Moreno-Azanza, M., & Canudo, J. I. The dinosaur assemblage of the El Castellar Formation
(Upper Hauterivian - Lowermost Barremian, Teruel, Spain)
Gasulla, J.M., Sanz, J.L., Ortega, F., & Escaso, F. Iguanodon bernissartensis from the Early Aptian of Morella
(Castellón, Spain)
Godefroit, P., Golovneva, L., Shchepetov, S. & Alexeev, P. The last polar dinosaurs: high diversity of latest
Cretaceous arctic dinosaurs in Russia
Gregorova, R. Remoras (Teleostei, Echeneidae) from the Oligocene of the West Carpathians (Czech Republic)
Hendrickx, C., Brusatte, S., Young, M., Rayfield, E., Ruta, M. & Barrett, P. Diversity and disparity in sauropod
dinosaurs
Hendrickx, C. & Buffetaut, E. Morphofunctional analysis of spinosaurid quadrates
Krempaská, Z. Dragons (Ursus spelaeus) by Johan Paterson Hain (1615 -1675) from Pieniny (Slovakia)
Métais, G., Qi, T., Guo, J. & Beard, C. The basal radiation of Ruminantia in Asia and the "Dichobunoid Gordian
Knot"
Pouech, J. & Mazin, J.-M. Description and West European affinities of the mammalian fauna of Cherves-de-
Cognac
Robaszynski, F. Foraminifera from the Bernissart boring F3 in the context of the Mid-Cretaceous transgression
Santos-Cubedo, A., de Santisteban, C. & Galobart, A. New dinosaur findings from Arcillas de Morella
Formation (Spain)
Spagna, P., Dupuis, C. & Yans, J. Comparison of sedimentological data in two wealden facies sites: The
Bernissart natural pit and the Danube-Bouchon Quarry of Hautrage (Mons Basin, Belgium). Implication on
their geodynamic history
Valentin, X., Godefroit, P., Tabuce, R., Vianey-Liaud, M., Garcia, G. & Wu, W. First Maastrichtian vertebrate
assemblage from Provence (Vitrolles-La-Plaine, France)
Vincent, P. & Suan, G. Diversity and paleogeographic distribution of Early Jurassic plesiosaurs
Wu, W., Dong, Z., Sun, Y., Li, C. & Li, T. A sauropod dinosaur from the Cretaceous of Jiutai, Jilin, China

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
FISH FAUNA FROM OLIGOCENE BITUMINOUS MARLS OF MARGINAL FOLDS NAPPE,
EAST CARPATHIANS, ROMANIA
Dorin Sorin BACIU, Crina MICLU , Ionu GRDIANU, Mihai NICULIl .............................. 15

NEW EARLY CRETACEOUS MULTITUBERCULATE FOSSILS FROM THE IBERIAN
PENINSULA
Ainara BADIOLA, José Ignacio CANUDO, Gloria CUENCA-BESCÓS ........................................ 16

CRETACEOUS COASTAL ENVIRONMENTS IN THE MONS BASIN: EVIDENCES FROM
CENOMANIAN DEPOSITS IN THE BETTRECHIES QUARRY
Jean-Marc BAELE, Florence QUESNEL, Chantal BOURDILLON, Christian DUPUIS ................ 17

THE MARINE REPTILE FAUNAS FROM THE MAASTRICHTIAN (LATEST CRETACEOUS)
PHOSPHATES OF MOROCCO
Nathalie BARDET ............................................................................................................................. 18

OLIGO-MIOCENE ODONTOCETES FROM THE MALTESE ISLANDS: STRATIGRAPHICAL
DISTRIBUTION AND EVOLUTION
Giovanni BIANUCCI, Michael GATT, Rita CATANZARITI ......................................................... 19

PALAEOZOIC BIODIVERSITY, ECOSYSTEMS AND EVOLUTION: THE CASE OF
ORDOVICIAN TO DEVONIAN VERTEBRATE-DOMINATED ASSEMBLAGES
Alain BLIECK ................................................................................................................................... 20

EARLY CRETACEOUS DINOSAURS AND PTEROSAURS FROM THE EASTERN PARIS
BASIN
Eric BUFFETAUT ............................................................................................................................. 21

THE HISTORICAL SIGNIFICANCE OF THE IGUANODON DISCOVERIES AT BERNISSART
Eric BUFFETAUT ............................................................................................................................. 22

EARLY CRETACEOUS FLORAS FROM SIBERIA, MONGOLIA, AND NORTH-EASTERN
CHINA
Eugenia V. BUGDAEVA, Valentina S. MARKEVICH .................................................................... 23

TO BE OR NOT TO BE: CAN CONODONTS BE VERTEBRATES?
Pierre BULTYNCK ........................................................................................................................... 24

THE BRAINCASE OF RHABDODON (DINOSAURIA: ORNITHOPODA): NEW SPECIMENS
FROM THE LATE CRETACEOUS OF SOUTHERN FRANCE
Phornphen CHANTHASIT ............................................................................................................... 25

NECK UTILIZATION AND FEEDING STRATEGY IN SAUROPOD DINOSAURS
Andreas CHRISTIAN, Gordon DZEMSKI, Jan-Thomas MÖLLER ................................................ 26

WHAT'S NEW IN THE WORLD OF DEVONIAN TETRAPODS?
Jennifer A. CLACK ........................................................................................................................... 27

BIODIVERSITY OF THE DEVONIAN TETRAPOD-BEARING LOCALITY OF STRUD,
BELGIUM
Gaël CLEMENT, Cyrille PRESTIANNI .......................................................................................... 28

MAASTRICHTIAN LAND VERTEBRATES IN RUSCA MONTAN BASIN (ROMANIA)
Vlad CODREA, Pascal GODEFROIT, Thierry SMITH, Ctlin JIPA-MURZEA .......................... 29

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
SPECIALIZED DENTITION IN LOWER CRETACEOUS FRESHWATER HYBODONT SHARKS
Gilles CUNY, Lionel CAVIN, Varavudh SUTEETHORN ............................................................... 30

PALYNOLOGICAL OVERVIEW OF WEALDEN FACIES SEDIMENTS FROM BELGIUM AND
NORTHERN FRANCE
Jean DEJAX, Denise PONS, Johan YANS ........................................................................................ 31

FROM LAND TO SEA: THE EARLY EVOLUTION OF CETACEANS
Christian DE MUIZON, Jonathan GEISLER .................................................................................... 32

VERTEBRATE REMAINS IN THE 2003 BERNISSART DRILL: HISTOLOGICAL ASSESSMENT
Armand DE RICQLES, Pascal GODEFROIT, Johan YANS ........................................................... 33

IGUANODON-LIKE FOOTPRINTS FROM THE ENCISO GROUP (APTIAN, LOWER
CRETACEOUS) OF LA RIOJA (CAMEROS BASIN, SPAIN)
Ignacio DÍAZ-MARTÍNEZ, Félix PÉREZ-LORENTE, Xabier PEREDA-SUBERBIOLA, José
Ignacio CANUDO ............................................................................................................................... 34

TRANSMISSION ELECTRON MICROSCOPY OF SAUROPOD BONES: A LOOK INSIDE THE
NANOSTRUCTURE
Maïtena DUMONT, András BORBELY, Aleksander KOSTKA, Anke PYZALLA ........................ 35

PTEROSAUR FLIGHT DYNAMICS AND THE EVOLUTION OF FLIGHT STYLE WITHIN THE
LINEAGE: INITIAL EXPERIMENTAL FINDINGS
Ross ELGIN, Edina PRONDVAI, Eberhard FREY, Wolf KRÜGER .............................................. 36

DIET EVIDENCES ON VISEAN CRUSHING-TOOTH (COCHLIODUS) FROM SOIGNIES
(BELGIUM)
Fabrice FACK, Claire DERYCKE ..................................................................................................... 37

PERMOCARBONIFEROUS AMNIOTES FROM FRANCE: A STATE OF THE ART AND
PERSPECTIVE
Jocelyn FALCONNET ...................................................................................................................... 38

NEW DATA ON THE PALAEOBIOGEOGRAPHY OF TOARCIAN (LOWER JURASSIC)
ICHTHYOSAURS
Valentin FISCHER, Myette GUIOMAR, Pascal GODEFROIT ....................................................... 39

A NEW IGUANODON SPECIES (DINOSAURIA, ORNITHISCHIA) FROM THE LOWER
CRETACEOUS OF SORIA (SPAIN)
Carolina FUENTES VIDARTE, Manuel MEIJIDE CALVO, Federico MEIJIDE FUENTES, Manuel
MEIJIDE FUENTES ........................................................................................................................... 40

CHELONIANS FROM THE LATE MIOCENE OF CHAD (THE HOMINID SITE TM 266)
Géraldine GARCIA, Likius ANDOSSA, Hassan-Taisso MACKAYE, Patrick VIGNAUD, Michel
BRUNET ............................................................................................................................................. 41

AMNIOTIC EGGSHELLS FROM THE HAlEG BASIN (UPPER CRETACEOUS, ROMANIA)
Géraldine GARCIA, Pascal GODEFROIT, Thierry SMITH, Jimmy VAN ITTERBEECK, Xavier
VALENTIN, Vlad CODREA ............................................................................................................. 42

THE DINOSAUR ASSEMBLAGE OF THE EL CASTELLAR FORMATION (UPPER
HAUTERIVIAN LOWERMOST BARREMIAN, TERUEL, SPAIN)
José Manuel GASCA, Miguel MORENO-AZANZA, José Ignacio CANUDO ................................ 43

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
IGUANODON BERNISSARTENSIS FROM THE EARLY APTIAN OF MORELLA (CASTELLÓN,
SPAIN)
José Miguel GASULLA, José Luís SANZ, Francisco ORTEGA, Fernando ESCASO .................... 44

THE DOMESTICATION OF THE DOG DURING THE UPPER PALAEOLITHIC
Mietje GERMONPRÉ ....................................................................................................................... 45

EARLY LAND PLANT EVOLUTION: A TREMENDOUS SUCCESS STORY
Philippe GERRIENNE ...................................................................................................................... 46

EARLY MAMMALIAN BRAIN DIVERSITY: INSIGHTS FROM MULTITUBERCULATES
Emmanuel GILISSEN, Thierry SMITH ............................................................................................ 47

THE LAST POLAR DINOSAURS: HIGH DIVERSITY OF LATEST CRETACEOUS ARCTIC
DINOSAURS IN RUSSIA
Pascal GODEFROIT, Lina GOLOVNEVA, Sergei SCHEPETOV, Géraldine GARCIA, Pavel
ALEKSEEV ........................................................................................................................................ 48

A NEW `MIDDLE' CRETACEOUS VERTEBRATE LOCALITY FROM JILIN PROVINCE,
NORTH-EASTERN CHINA
Pascal GODEFROIT, JIN Liyong, CHEN Jun, Richard BUTLER .................................................. 49

FROM PAST COLLECTIONS OF BERNISSART TO RECENT COLLECTIONS OF HAUTRAGE:
NEW TAXONOMICAL AND ENVIRONMENTAL INSIGHTS
Bernard GOMEZ, Johan YANS, Thomas GILLOT, Paul SPAGNA, Clément COIFFARD, Véronique
DAVIERO-GOMEZ ........................................................................................................................... 50

ZALMOXES, RHABDODON AND RHABDODONTIDAE: HOW MANY GENERA AND HOW
MANY SPECIES?
Florent GOUSSARD .......................................................................................................................... 51

REMORAS (TELEOSTEI, ECHENEIDAE) FROM THE OLIGOCENE OF THE WEST
CARPATHIANS (CZECH REPUBLIC)
Ruzena GREGOROVA ..................................................................................................................... 52

DESCRIPTION OF THE MASTICATORY APPARATUS OF LAONASTES AENIGMAMUS
(RODENTIA, DIATOMYIDAE), NEW INSIGHTS INTO THE EVOLUTION OF
HYSTRICOGNATHY
Lionel HAUTIER, Soonchan SAKSIRI ............................................................................................ 53

DIVERSITY AND DISPARITY IN SAUROPOD DINOSAURS
Christophe HENDRICKX, Steve BRUSATTE, Mark YOUNG, Emily RAYFIELD, Marcello RUTA,
Paul BARRETT .................................................................................................................................. 54

MORPHOFUNCTIONAL ANALYSIS OF SPINOSAURID QUADRATES
Christophe HENDRICKX, Eric BUFFETAUT ................................................................................ 55

CONTRIBUTION TO THE EVOLUTIONARY HISTORY OF THE BEAKED WHALES
Ella HOCH ......................................................................................................................................... 56

DRAGONS (URSUS SPELAEUS) BY JOHAN PATERSON HAIN (1615 -1675) FROM PIENINY
(SLOVAKIA)
Zuzana KREMPASKÁ ...................................................................................................................... 57

NEW STEM SPERM WHALES FROM THE MIOCENE OF PERU

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
Olivier LAMBERT, Giovanni BIANUCCI, Christian de MUIZON, Alton C. DOOLEY, Jr .......... 58

IGUANODON'S BRAIN AND PERSPECTIVES ON ORNITHOPOD EVOLUTION
Pascaline LAUTERS, Walter COUDYZER, Martine VERCAUTEREN, Pascal GODEFROIT ..... 59

DIAGENESIS OF THE FOSSIL BONES OF IGUANODON BERNISSARTENSIS
Thierry LEDUC .................................................................................................................................. 60

INSULARITY AND DWARFISM IN LATE CRETACEOUS EUROPEAN DINOSAURS
Jean LE LOEUFF .............................................................................................................................. 61

MISSISSIPIAN FORMATIONS AND CRETACEOUS TECTONICS: THE IMPACT OF THE
CURRENT GEOTHERMAL RESERVOIR GENESIS ON THE PAST LANDSCAPES OF THE
MONS BASIN
Luciane LICOUR ............................................................................................................................... 62

REARING FOR FOOD? KINETIC/DYNAMIC MODELING OF BIPEDAL/TRIPODAL POSES IN
SAUROPOD DINOSAURS
Heinrich MALLISON ........................................................................................................................ 63

A REEVALUATION OF THE EVIDENCE SUPPORTING AN UNORTHODOX HYPOTHESIS ON
THE ORIGIN OF EXTANT AMPHIBIANS
David MARJANOVI, Michel LAURIN ......................................................................................... 64

VEGETATION FROM LATE CRETACEOUS DINOSAUR ECOSYSTEMS IN AMUR RIVER
REGION (RUSSIA AND CHINA)
Valentina MARKEVICH, Eugenia BUGDAEVA, Yury BOLOTSKY ........................................... 65

3D MODELLING OF THE PALEOZOIC TOP SURFACE IN THE BERNISSART AREA AND
INTEGRATION OF DATA FROM BOREHOLES DRILLED IN THE "CRAN AUX
IGUANODONS"
Thierry MARTIN, Johan YANS, Christian DUPUIS, Paul SPAGNA, Olivier KAUFMANN ........ 66

STRATIGRAPHY OF THE HAINE GROUP ("MEULE" SEDIMENTS) OVERLYING THE
WEALDEN FACIES IN THE BER 3 BOREHOLE (BERNISSART, MONS BASIN, BELGIUM)
Edwige MASURE, Johan YANS ....................................................................................................... 67

COMPOSITION AND BIOGEOGRAPHIC AFFINITIES OF THE MIDDLE EOCENE MESSEL
AVIFAUNA
Gerald MAYR .................................................................................................................................... 68

THE BERNISSARTIDS: COMMON EUROPEAN CRUSHING CROCODILES
Jean-Michel MAZIN, Joane POUECH, Julien RASLAN-LOUBATIE ............................................ 69

THE BASAL RADIATION OF RUMINANTIA IN ASIA AND THE "DICHOBUNOID GORDIAN
KNOT"
Grégoire MÉTAIS, Tao QI, Jianwei GUO, Chris BEARD ............................................................... 70

CARBONIFEROUS AMPHIBIANS AND REPTILES EXPERIMENTATION AND
DIVERSIFICATION
Andrew R. MILNER .......................................................................................................................... 71

NEW DATA ON THE VALANGINIAN-HAUTERIVIAN REPTILE OOTAXA OF THE IBERIAN
RANGE (NE OF SPAIN)
Miguel MORENO-AZANZA, José Manuel GASCA, José Ignacio CANUDO ............................... 72

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009

PALAEOENVIRONMENTAL CONTEXT OF THE `BLACK MARBLE' OF DENÉE (VISÉAN,
BELGIUM)
Bernard MOTTEQUIN, Edouard POTY .......................................................................................... 73

IGUANODONTIANS FROM THE WEALDEN OF BRITAIN AND EUROPE
David B. NORMAN ........................................................................................................................... 74

A NEW SAUDI ARABIAN TAXON OF ACANTHOTHORACI (PLACODERMI): QUESTIONS
ON PLATE HOMOLOGIES AND GROWTH
Sébastien OLIVE, Daniel GOUJET, Hervé LELIEVRE ................................................................... 75

THE LATE CRETACEOUS CONTINENTAL VERTEBRATE FAUNA FROM IHARKÚT,
WESTERN HUNGARY A REVIEW OF THE NEWEST RESULTS
Attila SI ............................................................................................................................................ 76

A NEW IGUANODONTIAN DINOSAUR FROM THE EARLY CRETACEOUS YIXIAN
FORMATION OF WESTERN LIAONING, CHINA
Rui PAN, Yihan WANG, Xiaolin WANG ......................................................................................... 77

LOWER CRETACEOUS DINOSAURS OF SPAIN: AN OVERVIEW BASED ON SKELETAL
REMAINS
Xabier PEREDA SUBERBIOLA, José Ignacio RUIZ-OMEÑACA, José Ignacio CANUDO, Fidel
TORCIDA, José Luis SANZ .............................................................................................................. 78

EARLY CRETACEOUS ORNIHTOPOD DINOSAURS FROM ROMANIA
Erika POSMOSANU ......................................................................................................................... 79

FAUNAL ASSEMBLAGES AT THE JURASSIC/CRETACEOUS BOUNDARY: COMPARISON
BETWEEN SOME WEST EUROPEAN CONTINENTAL SITES
Joane POUECH, Jean-Michel MAZIN .............................................................................................. 80

DESCRIPTION AND WEST EUROPEAN AFFINITIES OF THE MAMMALIAN FAUNA OF
CHERVES-DE-COGNAC
Joane POUECH, Jean-Michel MAZIN .............................................................................................. 81

THE KARSTIC PHENOMENON OF BERNISSART PIT AND THE GEOMORPHOLOGIC
SITUATION IN THE MESOZOIC TIMES
Yves QUINIF ..................................................................................................................................... 82

PRESENCE OF SEBECOSUCHIAN CROCODYLIFORMS IN THE LATE CRETACEOUS OF
EUROPE
Márton RABI ...................................................................................................................................... 83

FORAMINIFERA FROM THE BERNISSART BORING F3 IN THE CONTEXT OF THE MID-
CRETACEOUS TRANSGRESSION
Francis ROBASZYNSKI ................................................................................................................... 84

THE ATTEMPTED THEFT OF DINOSAUR SKELETONS DURING THE GERMAN
OCCUPATION OF BELGIUM 1914-1918 AS A TOPIC OF THE HISTORY OF SCIENCE AND
THE HISTORY OF TOTAL WAR
Christoph ROOLF .............................................................................................................................. 85

THE RECORD OF CLIMATE AND EUSTATIC CHANGES DURING THE LOWER
CRETACEOUS IN THE "ARCILLAS DE MORELLA FORMATION" (SPAIN)

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
Carlos de SANTISTEBAN, Andrés SANTOS-CUBEDO ................................................................ 86

NEW DINOSAUR FINDINGS FROM ARCILLAS DE MORELLA FORMATION (SPAIN)
Andrés SANTOS-CUBEDO, Carlos de SANTISTEBAN, Angel GALOBART .............................. 87

ORGANIC MATTER CHARACTERIZATION AND ORGANIC CARBON ISOTOPES RECORD
IN THE EARLY CRETACEOUS LACUSTRINE SETTING OF BERNISSART (BELGIUM)
Johann SCHNYDER, Jean DEJAX, Eddy KEPPENS, Thanh Thuy NGUYEN TU, Paul SPAGNA,
Armelle RIBOULLEAU, Johan YANS .............................................................................................. 88

MAMMALIAN FAUNA OF HYENA DEN CAVE (ALTAI, RUSSIA)
Natalia SERDYUK, Sergei VASILIEV ............................................................................................. 89

COMPARISON OF SEDIMENTOLOGICAL DATA IN TWO WEALDEN FACIES SITES: THE
BERNISSART NATURAL PIT AND THE DANUBE-BOUCHON QUARRY OF HAUTRAGE
(MONS BASIN, BELGIUM). IMPLICATION ON THEIR GEODYNAMIC HISTORY
Paul SPAGNA, Christian DUPUIS, Johan YANS ............................................................................. 90

THE PROBLEMATIC FOSSIL BALEEN WHALES FROM BELGIUM
Mette E. STEEMAN .......................................................................................................................... 91

QUANTIFYING GROWTH RATES IN ISLAND DWARF SAUROPODS
Koen STEIN, Martin SANDER ......................................................................................................... 92

THE AFRICAN COUSINS OF THE EUROPEAN IGUANODONS
Philippe TAQUET ............................................................................................................................. 93

FIRST MAASTRICHTIAN VERTEBRATE ASSEMBLAGE FROM PROVENCE (VITROLLES-
LA-PLAINE, FRANCE)
Xavier VALENTIN, Pascal GODEFROIT, Rodolphe TABUCE, Monique VIANEY-LIAUD,
Géraldine GARCIA, Wenhao WU
.............................................................................................................................................................. 94

CRUSTAL TECTONIC CONTROL OF THE EARLY CRETACEOUS DEPOSITS IN THE MONS
BASIN (BELGIUM)
Sara VANDYCKE, Paul SPAGNA ................................................................................................... 95

ORIGINE DE L'HOMME: LES NOUVELLES DU TCHAD
Patrick VIGNAUD, MPFT ................................................................................................................ 96

DIVERSITY AND PALEOGEOGRAPHIC DISTRIBUTION OF EARLY JURASSIC
PLESIOSAURS
Peggy VINCENT, Guillaume SUAN ................................................................................................ 97

THE TERRESTRIAL LOWER CRETACEOUS OF GERMANY ("WEALDEN")
Volker WILDE, Walter RIEGEL ....................................................................................................... 98

A SAUROPOD DINOSAUR FROM THE CRETACEOUS OF JIUTAI, JILIN, CHINA
Wen-hao WU, Zhi-ming DONG, Yue-wu SUN, Chun-tian LI, Tao LI ............................................. 99

INTEGRATED RESULTS OF THE NEW MATERIAL FROM THE 2002-2003 BOREHOLES IN
BERNISSART (MONS BASIN, BELGIUM)
Johan YANS, Bernard ANDREU, Jean-Marc BAELE, Colette CORNET, Armand DE RICQLES,
Jean DEJAX, Christian DUPUIS, Bruno GALBRUN, Thomas GERARDS, Philippe GERRIENNE,
Pascal GODEFROIT, Bernard GOMEZ, René GOSSELIN, Thierry LEDUC, Gilles PETIT, Denise

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
PONS, Alain PREAT, Francis ROBASZYNSKI, Johann SCHNYDER, Thierry SMITH, Paul
SPAGNA, Etienne STEURBAUT, Louis TAVERNE, Jean-Pierre TSHIBANGU, Jimmy VAN
ITTERBEECK, Sara VANDYCKE, Céline VANNESTE ................................................................. 100

CARBON ISOTOPES ON WOODY MATERIAL FROM WEALDEN FACIES OF HAUTRAGE
(MONS BASIN, BELGIUM)
Johan YANS, Jean DEJAX, Thomas GERARDS, Philippe GERRIENNE, Paul SPAGNA, Eddy
KEPPENS............................................................................................................................................ 101

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
FISH FAUNA FROM OLIGOCENE BITUMINOUS MARLS OF
MARGINAL FOLDS NAPPE, EAST CARPATHIANS, ROMANIA

Dorin Sorin BACIU 1, Crina MICLU 1, Ionu GRDIANU 2, Mihai NICULIl 3

1 Department of Geology, "Al. I. Cuza" University of Ia i (Romania), Bd. Carol I, 20A, 700505-Ia i
(Romania), dsbaciu@clicknet.ro, crina_miclaus@yahoo.co.uk
2 The Museum of Natural Sciences, str. Petru Rares no.26, 5600 Piatra Neamt, Romania,
igradianu@hotmail.com
3 Department of Geography, "Al. I. Cuza" University of Ia i (Romania), Bd. Carol I, 20A, 700505-Ia i
(Romania), mn_geogr@yahoo.com

The aims of this study are: 1) to establish the sedimentological significance of the Bituminous Marls,
which represent a lithostratigraphic marker of the Oligocene in East Carpathians, Romania, 2) to describe the
taxonomic composition of fish fossil species discovered in Bituminous Marls, and 3) to reconstruct the
palaeoecology and palaeobathimetry of fish assemblage based on ecology of recent fish fauna and
sedimentological conclusions.
In the East Carpathians, the Moldavide Nappe Complex represents a complicated tectonic edifice formed
by several tectonic units mainly consisting of Cretaceous to Early Miocene flysch and molasse deposits. The
Marginal Folds Nappe (or Vrancea Nappe) is one of the outermost units of the Moldavide Nappe Complex,
structurally interposed between the Tarcu and Subcarpathian Nappes respectively, and cropping out in several
tectonic half-windows.
During Eocene the sedimentation in area corresponding to MFN evolved from a calcareous one (Doamna
Limestones) toward a mainly siliciclastic one represented by a thick pelitic succession (Bisericani Formation) in
the late Eocene, grading into the well known Globigerina Marls at the Eocene-Oligocene boundary, and upward
into Oligocene deposits known in the Romanian geological literature as Lower Menilite, Bituminous Marl,
Lower Dyssodilic Shale with Kliwa Sandstones, Upper Dyssodilic Shale and Menilite, and Gura oimului
Formations.
The Bituminous Marls of Marginal Folds Nappe (or Vrancea Nappe) although seem to be a monotonous
deposit is, in fact rather a heterolithic deposit (mainly mudstone with sandstones and some conglomerates with
"green schist" clasts) consisting of many simple or composite bed-sets with different sedimentary structures (fine
lamination, hummocky and swaley-like cross stratification, large scale low angle cross stratification, wavy and
lenticular bedding) resulted from different processes from hemipelagic to tractive, and even debris flows. Beside
of sedimentary depositional structures, there are also deformational meta-depositional structures such as
convolute lamination in bituminous marls (s.s.), load casts and flame, and convolute lamination in sandstone
interlayers; load cast and flame structures are a result of a liquidization (liquefaction and/or fluidization) of
sandy sediments due to rapid sedimentation of sands over undercompacted muds while convolute lamination use
to appear because of current action, loading, sudden regular releases of pressure etc. Some post-depositional
structures can also be observed in Bituminous Marls, such as: dykes, sills and ptygmatic structures.
The existent facies, interpreted in terms of sedimentary processes, can be grouped in a facies association
which has many characteristics of a storm influenced mid- to outer shelf sedimentary environment/system. Some
intervals of wave influence can also be noticed.
A significant Oligocene fish fauna has been collected from Bituminous Marls of MFN in Bistria Half-
Window (Piatra Neam area). These fishes are well preserved and the collection contains specimens of more than
15 species representing about 11 families. The most important species include sardinas (Clupeidae), argentines
(Argentinidae), bristlemouth (Gonostomatidae), lightfishes (Photichthyidae), lanternfishes (Myctophidae), true
cods (Gadidae), boarfishes (Caproidae), seabasses (Serranidae), scabbardfishes (Trichiuridae), windowpanes
(Scophthalmidae). Well represented are mesopelagic species with luminescent organs (Gonostomatidae,
Photichthyidae, Myctophidae), but in the same time a lot of species prefer continental shelf, near bottom
(Argentinidae, Gadidae, Caproidae, Serranidae, Trichiuridae, Scophthalmidae).

Key words: fish fossils, Bituminous Marls, Oligocene, Marginal Folds Nappe (MFN), East Carpathians

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
NEW EARLY CRETACEOUS MULTITUBERCULATE FOSSILS FROM
THE IBERIAN PENINSULA

Ainara BADIOLA, José Ignacio CANUDO, Gloria CUENCA-BESCÓS

Universidad de Zaragoza, Ciencias de la Tierra (Área de Paleontología), Pedro Cerbuna, 12, 50009 Zaragoza,
Spain. Grupo Aragosaurus (http://www.aragosaurus.com), abadiola@unizar.es, jicanudo@unizar.es,
cuencag@unizar.es

The most abundant and diverse fossils of multituberculate mammals are from the Early Cretaceous of
Western Europe. They range from the Berriasian of Portugal and England up to the Barremian of Spain. Fossils
of Valanginian age are scarce, and no Hauterivian multituberculates in Europe is reported. The Early Cretaceous
multituberculate fossils from other areas of Laurasia and from Gondwana (Africa) are less diverse. New
multituberculate finds have recently been described in the late Hauterivian-early Barremian of Spain, in the La
Cantalera vertebrate locality, Teruel (Badiola et al., 2008), and in a Barremian or Aptian locality from the Tetori
Group of Japan (Kusuhashi, 2008). New Barremian specimens have also found in the Wessex Formation from
England (Sweetman, in press).
Here we report around sixteen new isolated teeth of the Valanginian/Hauterivian transition (the
Pochancalo site, Villanueva de Huerva Fm.), the Hauterivian/Barremian transition (the Cantalera site, Blesa
Fm.), and of the late Barremian (the Vallipón site, Artoles Fm.). These assemblages all come from the
Aragonese branch of the Iberian Ranges (north-eastern of Spain). This work summarizes these new finds and
updates the collection of isolated teeth (around twenty) from the Galve beds of Barremian age, previously
described by Crusafont and Gibert (1976). The new discoveries that we report here increase the resolution of the
multituberculate mammalian biostratigraphy and palaeobiogeography of these faunas during the Early
Cretaceous in the Iberian Peninsula: an upper tooth found in Pochancalo-1 is assigned to the Albionbaataridae
family, which represents the first early Hauterivian member in Europe and the first one in the Iberian Early
Cretaceous assemblages; a new pinheirodontid taxon, Cantalera abadi Badiola et al., 2008, has recently been
described in the La Cantalera site, together with the oldest representative of the eobaatarid Eobaatar Kielan-
Jaworowska et al., 1987 and a taxon described as Plagiaulacidae or Eobaataridae gen. et sp. indet. The latter is
also present in a site of Galve of the same age and probably belongs to a new taxon of the family Plagiaulacidae.
If this assignment is correct, it will be the first representative of this family in the Iberian Peninsula. In the late
Barremian site of Vallipón, high number of isolated lower and upper teeth of Eobaatar is found, together with
other pinheirodontid and paulchoffatiid specimens.
To date, the most complete Early Cretaceous multituberculate fossil record is present in the Iberian
Peninsula, which ranges from the Berriasian to late Barremian assemblages. These Iberian Early Cretaceous
multituberculate faunas comprise endemic Iberian Jurassic survivors (Paulchoffatiidae), those which are
currently only recorded in Britain and in the Iberian Peninsula (Pinheirodontidae, Albionbaataridae, and possibly
Plagiaulacidae), and others which are also registered in Asia (Eobaataridae). The presence of the eobaatarid
Eobaatar from the Hauterivian/Barremian transition to late Barremian in the Iberian Peninsula, and during the
Barremian in Britain and Aptian or Albian in Asia, suggest that geographical connections between these areas
could have existed either sporadically or constantly for most of the Early Cretaceous. This hypothesis is
supported by other palaeontological data such as the Gobiconodontidae mammals (Cuenca-Bescós and Canudo,
2003), some Sauropoda and Ornithopoda dinosaurs (Canudo et al., 2002).

Key words: multituberculates, Early Cretaceous, Iberian Peninsula, systematic, palaeobiogeography

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
CRETACEOUS COASTAL ENVIRONMENTS IN THE MONS BASIN:
EVIDENCES FROM CENOMANIAN DEPOSITS IN THE
BETTRECHIES QUARRY

Jean-Marc BAELE 1, Florence QUESNEL 2, Chantal BOURDILLON 3,
Christian DUPUIS 1

1 Faculté Polytechnique de Mons, rue de Houdain, 9, 7000 Mons, Belgique, jean-marc.baele@fpms.ac.be
2 BRGM, CDG/MA, 45060 Orléans, Bp6009, France, f.quesnel@brgm.fr
3 ERADATA, allée des magnolias, 5, 72100, Le mans, France, bourdillon@eradata.fr

Cenomanian marine transgressions in the Mons Basin left mixed siliclastics-carbonate deposits that are
well exposed in the Bettrechies quarry, Northern France (Robaszynski, 1975). Though they are not the oldest
remains of sea invasion pulses since "Wealden times" (Marlière, 1970), these deposits are overlying continental,
i.e. lignitic and/or pyritic sands and clays filling karsts in the Devonian bedrock. No dating is available yet for
these continental sediments but their lithology and overall stratigraphic location is similar to well established
Wealden facies in adjacent areas (i.e. Hautrage, Bernissart, etc.).
Investigations focused on the first, m-thick calcareous marine beds referred to as "Sarrazin" in local
denomination. The global succession is as follow: basal conglomerates, up to 1m thick, are overlaid by
calcarenites, up to 3m thick, showing one or two highly cemented beds, especially on top formation.

Conglomerates mainly consist of Lower Devonian sandstones pebbles in clayey/calcareous matrix that
deposited in littoral environment. They only occur in karst fillings where collapse of the underlying "Wealden"
sediments preserved them against erosion. Outside karstic areas, red cm-thick stromatolites are found instead, as
discontinuous layer on the Devonian bedrock. Their unusual morphology and growth pattern indicate that the
karsts induced ponds, possibly freshwater springs, in very shallow coastal environment. Together with
sedimentological evidences, this suggests a coastal landscape first dominated by highs, possibly cliffs, for the
emplacement of early Cenomanian deposits.
The overlying calcarenite shows significant lateral variability controlled by shallow marine dynamics
and bedrock topography. Petrographic evidences from stained thin-sections and cathodoluminescence analysis
show that lithification initiated in the marine vadose zone, suggesting that the hardened calcarenite beds could be
similar to some modern beachrocks. Asymmetrical and meniscus morphologies of the earliest spar cement along
with solution features are the strongest indications for such an environment. The cementation sequence actually
consists of syntaxial overgrowth alternating with carbonate mud infiltration related to (tidal?) sea level
fluctuations. Iron-stained shrub- or Frutexites-like structures (Flügel, 2004) occur in the mud filling, now
lithified micrite. They are indicative of bacterial activity, possibly by iron-oxidizing consortia similar to that
forming the underlying red stromatolithes, but scattered through the carbonate sediment. The role of bacterial
activity in the mud lithification is still unclear but is thought to be crucial in the early cementation of modern
beachrocks (Neumeier, 1998).
Thin, discontinuous stromatolithe crusts were also observed capping a gravel bed (storm deposit?)
intercalated in the calcarenites. They occur in dm-wide fossil "pools" that could be interpreted as animal
(dinosaur?) footprints.
This study, which is the first detailing the coastal depositional environment for the Cenomanian sediments
in the investigated area, opens perspectives in reconstructing the landscape evolution sequence from continental
("Wealden") to marine environments through Cretaceous times.

FLÜGEL, E., 2004. Microfacies of carbonate rocks. Analysis, interpretation and application., Springer, Berlin, 976 p.
MARLIERE, R., 1970. Géologie du bassin de Mons et du Hainaut : Un siècle d'histoire. Ann. Soc. Géol. Nord., 40 : 171-189.
NEUMEIER, U., 1998. Le rôle de l'activité microbienne dans la cimentation précoce des beachrocks (sédiments intertidaux). Terre et
Environnement, 12 : 1-183.
ROBASZYNSKI, F., 1975, In: Marlière, R. & Robaszynski, F., Document n° 9. Crétacé. Conseil Géologique - Commissions Nationales de
Stratigraphie, Ministère des Affaires Économiques, 53 p.

Key words: carbonate, diagenesis, coastal environment, Cretaceous, Mons Basin

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Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE MARINE REPTILE FAUNAS FROM THE MAASTRICHTIAN
(LATEST CRETACEOUS) PHOSPHATES OF MOROCCO

Nathalie BARDET

UMR 5143 du CNRS / Département Histoire de la Terre, Muséum National d'Histoire Naturelle, CP 38, 8 rue
Buffon, 75005 PARIS, France, bardet@mnhn.fr

The phosphates of Morocco (Maastrichtian-Lutetian) form part of the Mediterranean Tethyan
phosphogenic province - a complex of warm and shallow marine platforms characterised by intense phosphatic
sedimentation - that bordered the southern Tethys margin during the Late Cretaceous and Early Palaeogene.
These phosphates are very rich in marine vertebrate remains, especially selachians, bony fishes and reptiles that
are known since the pioneer work of Arambourg (1952). Since a decade, dynamised by the discovery of the early
proboscidean Phosphaterium (Gheerbrant et el., 1996), important fieldworks have been made in the framework
of and active french-moroccan. These resulted in the discovery of major marine and continental vertebrate
faunas, ranging from the Maastrichtian to the Ypresian, and including representatives of selachians, bony fishes,
reptiles (including birds) and mammals (see Jalil et al., 2006).
The Late Cretaceous phosphatic series of Morocco are early to late Maastrichtian in age. They have yielded one
of the richest and most diversified marine reptile assemblages of the world. The squamates, and more especially
the large and fully adapted to marine life Mosasauridae, are by far the most abundant and diversified, including
six coeval genera of the clades halisaurine (Halisaurus), mosasaurine (Mosasaurus, Prognathodon, Globidens,
Carinodens) and russellosaurine ('Platecarpus') (Bardet et al., 2008). They also include the poorly known
varanoid Pachyvaranus. The plesiosaurs are less diversified than the mosasaurids and represented up to now by a
new elasmosaurid genus only. The crocodilians are very scarce in the Maastrichtian levels and only known by
the gavialoid eusuchian Ocepesuchus. Whereas turtles are highly diversified in the Palaeogene and represented
by both bothremydid pleurodirans and chelonioid cryptodirans, only scarce chelonioid remains have been
described up to now in the Maastrichtian phosphates. In addition to these marine reptile faunas, scarce
continental remains of azhdarchid pterosaurs as well as of theropod and titanosauriform dinosaurs have been
found in these Maastrichtian levels.

The continuous Late Cretaceous - Early Palaeogene phosphatic series of Morocco provide and
exceptional framework for an in situ study of the K/T faunal turnover, as exemplified by the extinction of
mosasaurs, plesiosaurs, dinosaurs and pterosaurs at the end of the Cretaceous and the rapid radiation of
crocodiles, chelonians, birds and mammals in the Early Palaeogene. As far as marine reptiles are concerned, the
mosasaurid squamates were highly diversified and abundant during all the Maastrichtian stage and disappeared
near the K/T boundary, whereas the crocodyliforms remained very scarce at this time. Mirroring this, eusuchian
and dyrosaurid crocodyliforms highly diversified as soon as the beginning of the Palaeogene (Jouve et al., 2008).
On a palaeobiogeographical point of view, these Maastrichtian marine reptile assemblages are characteristic of
the Southern Tethys margin (between palaeolatitudes 20° N-20°S) and differs from the contemporaneous ones of
the Northern Tethys Margin (around palaeolatitude 35° N) (Bardet, 2004). These observed paleoecological
segregation is interpreted as revealing different faunal subprovinces, probably linked to palaeolatitudinal
gradients.

ARAMBOURG, C., 1952. Les vertébrés fossiles des gisements de phosphates (Maroc-Algérie-Tunisie). Notes et Mémoires du Service
Géologique du Maroc, 92: 1-372.
BARDET, N., 2004. Les faunes de reptiles marins des marges Nord et Sud de la Téthys méditerranéenne durant le Crétacé supérieur:
systématique et paléobiogéographie. Habilitation à Diriger des Recherches, Université Paris VI, Paris, 95 p.
BARDET, N., PEREDA SUBERBIOLA, X., SCHULP, A., & BOUYA, B. 2008a. New material of Carinodens (Squamata, Mosasauridae)
from the Maastrichtian (Late Cretaceous) Phosphates of Morocco. Bulletin of the Fort Hays State University, Special Issue,3: 29-
36.
GHEERBRANT, E., SUDRE, J. & CAPPETTA, H. 1996. A Palaeocene proboscidean from Morocco. Nature, 383: 68-71.
JALIL, N.-E., BARDET, N., BOURDON, E., CAPPETTA, H., GHEERBRANT, E., JOUVE, S., NOUBHANI, A., OUANAIMI, F.,
PEREDA SUBERBIOLA, X., VINCENT, P., AMAGHZAZ, M., BOUYA, B., & MCHICHI, M. 2006. Les vertébrés fossiles des
phosphates du Maroc. Nouvelle convention et état des connaissances. COVAPHOS II, Marrakech, p. 01-04-19.
JOUVE, S., BARDET, N., JALIL, N.E., PEREDA SUBERBIOLA, X., BOUYA, B., & AMAGHZAZ M., 2008. The oldest African
crocodylian: phylogeny, palaeobiogeography, and differential survivorship of marine reptiles through the Cretaceous-Tertiary
boundary. Journal of Vertebrate Paleontology, 28(2): 409-421.

Key words: marine reptiles, phosphates, Morocco, K/T boundary, palaeobiogeography

18

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
OLIGO-MIOCENE ODONTOCETES FROM THE MALTESE
ISLANDS: STRATIGRAPHICAL DISTRIBUTION AND
EVOLUTION

Giovanni BIANUCCI 1, Michael GATT 2, Rita CATANZARITI 3

1 Dipartimento di Scienze della Terra, Università di Pisa, via S. Maria, 53, I-56126, Pisa (Italy),
bianucci@dst.unipi.it
2Cidaris", 18, Hal-Bajjada Street, Rabat RBT2030 (Malta), cidaris@waldonet.net.mt
3 CNR - Istituto di Geoscienze e Georisorse, via S. Maria, 53, I-56126, Pisa (Italy), catanzariti@igg.cnr.it

Outcropping in the Maltese archipelago is a unique marine sedimentary succession punctuated by
several hiatuses and deposited over a wide age interval, spanning from the Late Oligocene to the Late Miocene.

Fossil whales have been reported from these sediments as early as the 17th century, where in 1670 the
Italian Agostino Scilla described and illustrated for the first time in the history of science, a fossil whale in his
famous work "La vana speculazione disingannata dal senso". More recently published records are very scarce
and prevalently limited to poorly diagnostic material.

In recent years, new odontocete fossil remains have been discovered in almost all the outcropping
stratigraphical sequence, and more or less in continuity. Although this new material is rather fragmentary
(prevalently ear bones and teeth), in most cases, the identification to family level was possible. A preliminary
analysis of these fossils and a revision of the previously published records permitted the identification of at least
seven odontocete families. Three of them (Kogiidae, Pontoporiidae and Waipatiidae) constitute first records for
the Mediterranean.

The exact stratigraphical provenance of most of this new material is known. Furthermore, calcareous
nannoplancton analysis from sediments in association with some of the studied material, made possible their
localisation within the sedimentary succession. As a result we were able to recognize the following general trend
and changes in the faunal composition:

1. In the older strata, particularly the Lower and Middle members and in the Lower Main Phosphorite
Conglomerate Bed of the Globigerina Limestone Formation (Chattian-Burdigalian), the fossil record is
prevalently represented by Platanistoidea with Waipatiidae and Squalodontidae. The waipatiid record
reflects the Mediterranean-Indopacific distribution of this family possibly favored by the wide direct
communications existing between these waters during the Late Oligocene-Early Miocene.
2. In the Upper Main Phosphorite Conglomerate Bed and in the Upper member of the Globigerina Limestone
Formation (Langhian) the odontocete fauna is represented by Waipatiidae, stem Physeteroidea, Physeteridae
and Eurhinodelphinidae.
3. In the Blue Clay Formation (Serravallian and/or late Tortonian) the fossil assemblage is represented by stem
Physeteroidea, Ziphiidae, Kentriodontidae and Pontoporiidae. The pontopriid record confirms the wide
geographical distribution of these dolphins, today represented only by Pontoporia from the eastern south-
American coasts.
4. In the Greensand and Upper Coralline Formations (late Tortonian and/or Messinian) stem Physeteroidea,
Kogiidae, Ziphiidae and possible Pontoporiidae and other indeterminate Delphinida are reported. Although
the Late Miocene record from Malta is rather fragmentary, none of the new fossil material can be
unquestionably referred to Delphinidae, confirming that this family appears only in the Pliocene, at least in
the Mediterranean.

This general trend, in part reflecting a similar tendency observed in the whole Mediterranean, could be
related to a global evolution of the whale fauna and secondarily to more local factors due to the environmental
changes in the depositional basin.

Key words: Cetacea, Odontoceti, Malta, Oligocene, Miocene

19

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
PALAEOZOIC BIODIVERSITY, ECOSYSTEMS AND EVOLUTION:
THE CASE OF ORDOVICIAN TO DEVONIAN
VERTEBRATE-DOMINATED ASSEMBLAGES

Alain BLIECK

Université de Lille 1 : Sciences de la Terre, UMR 8157 du CNRS « Géosystèmes » (LP3), F-59655 Villeneuve
d'Ascq cedex (France), Alain.Blieck@univ-lille1.fr

Vertebrates, excluding conodonts as defined by e.g. Janvier (1996), form a monophyletic taxon of
chordate animals known possibly from the Early Cambrian and certainly from the Early Ordovician to Present.
They encountered major steps of evolution during the Palaeozoic, and in particular during the Early (Cambrian-
Ordovician) and Middle (Silurian-Devonian) Palaeozoic. The first fossilized creature possibly attributed to
vertebrates is Haikouichthys (Myllokunmingia) from the Early Cambrian Konservat-Fossil-Lagerstätte of
Chengjiang, China (e.g. Shu et al. 2003). Some 40 Ma later, vertebrates made their first adaptive radiation during
the Ordovician, for which we recognize two major palaeobiogeographic provinces called the Gondwana
Endemic Assemblage and the Laurentia-Baltica-Siberia Assemblage (Blieck & Turner 2003). Ordovician
vertebrates come from mostly siliciclastic series with trace fossils and lingulid brachiopods, interpreted as
transitional to coastal marine environments (e.g. Allulee & Holland 2005, Davies et al. 2007). After the
Hirnantian glaciation, which does not seem to have been an actual extinction event for vertebrates, the group
realized a new radiation during the Silurian and Devonian (Long 1993). Silurian vertebrate assemblages are, as
the Ordovician ones, dominated by agnathan forms, the ostracoderms. Silurian vertebrates from the East Baltic
basin have been collected in all the environments of the marine epicontinental platform up to lagoonal
environments (e.g. Märss & Ejnasto 1978, Schultze 1999). Early Devonian vertebrate assemblages are also
dominated by agnathans, but many jawed forms, the Gnathostomes, began to become more and more abundant,
diversified and dominant upon agnathans at that time when placoderms was the most important group of
Gnathostomes (Blieck & Janvier 1999). During the Middle and Late Devonian, most vertebrate assemblages are
dominated by Gnathostomes such as placoderms, acanthodians, chondrichthyans, sarcopterygians. Among the
latter, tetrapods appeared in the Late Devonian and are now known from the Old Red Sandstone Continent,
North China and eastern Gondwana (Autralia) (Clack 2002, Clément et al. 2004). Devonian vertebrates are both
known from continental to near-shore marine, Old Red Sandstones (ORS) and associated facies, and from
marine facies where they are often collected with conodonts and many other invertebrates. In those various
environments they developed many different convergent adaptations, and in particular numerous spiny
phenotypes in the ORS (such as long rostra, lateral expansions as the cornual plates or cornua of ostracoderms,
dorsal spines, etc.: Janvier 1996). At the transition from Devonian to Carboniferous, another major event
occurred, viz. the ecological replacement of placoderms by chondrichthyans (Long 1990). After the extinction of
ostracoderms at the Frasnian-Famennian boundary, and the extinction of placoderms at the Devonian-
Carboniferous boundary, chondrichthyans and actinopterygians became dominant in the fish faunas. This is
particularly evident in the Namurian Konservat-Fossil-Lagerstätte of Bear Gulch, Montana, USA (Lund &
Grogan 2005). Klug et al. (in press), through a review of ecosystems from the latest Proterozoic to the Early
Carboniferous, recognized in the history of marine life a series of major steps among which the Devonian
Nekton Revolution. In fact, as concerned with vertebrates, their first explosive radiation is Silurian-Early
Devonian in age and mostly due to ostracoderms: this is the Nekton Revolution of vertebrates. Their second
main diversity peak is Late Devonian (Frasnian) in age and mostly due to Gnathostomes: this is the Predation
Revolution of vertebrates.

ALLULEE J.L. & HOLLAND S.M. 2005, Palaios 20: 518-533.
BLIECK A. & JANVIER P. 1999, in Boucot A.J. & Lawson J.D. eds, Paleocommunities ... Cambridge Univ. Press: 79-105.
BLIECK A. & TURNER S. 2003, Palaeogeogr., Palaeoclimat., Palaeoecol. 195: 37-54.
CLACK J.A. 2002, Gaining Ground ... Indiana Univ. Press, 369 p.
CLEMENT G. et al. 2004, Nature 427: 412-413.
DAVIES N.S. et al. 2007, Palaeogeogr., Palaeoclimat., Palaeoecol. 249: 18-35.
JANVIER P. 1996, Early Vertebrates, Oxford Sci. Publ., Clarendon Press, 393 p.
KLUG C. et al. in press, Geology.
LONG J.A. 1990, in McNamara K.J. ed., Evolutionary Trends, Belhaven Press: 255-278.
LONG J.A. ed. 1993, Palaeozoic vertebrate ... Belhaven Press, 369 p.
LUND R. & GROGAN E. 2005, Fossil fishes of Bear Gulch, http://www.sju.edu/research/bear_gulch/
MÄRSS T. & EJNASTO R. 1978, Eesti NSV Tead. Akad. Toimet. 27, K. Geol. 1: 16-22.
SHU D.-G. et al. 2003, Nature 421: 526-529.

Key words: vertebrate assemblages, palaeoenvironments, Old Red Sandstones, Nekton Revolution, extinctions

20

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
EARLY CRETACEOUS DINOSAURS AND PTEROSAURS FROM THE
EASTERN PARIS BASIN

Eric BUFFETAUT

Centre National de la Recherche Scientifique (UMR 8538), Laboratoire de Géologie de l'Ecole Normale
Supérieure, 24 rue Lhomond, 75231 Paris Cedex 05, France. eric.buffetaut@wanadoo.fr

The Early Cretaceous formations of the eastern Paris Basin (departments Aube, Meuse, Ardennes,
Haute-Marne), consisting mainly of shallow marine deposits, with some continental intercalations (notably of
Barremian age), have yielded relatively abundant tetrapod fossils, many of them found in the course of the
industrial exploitation of natural resources such as phosphate and building stone during the 19th century. Several
of these fossil-bearing formations are no longer easily accessible, and only a few new finds have been reported in
recent years. However, a number of hitherto unreported specimens have been found in old collections, which
need to be inspected in more detail. Besides marine forms such as ichthyosaurs, plesiosaurs, crocodilians and
turtles, dinosaurs and pterosaurs also occur, as shown in the following list.

Hauterivian:
Calcaire à Spatangues:

Ornithopoda: Iguanodon atherfieldensis

Pterosauria: Ornithocheiridae indet.
Barremian:

Ornithopoda: Iguanodon bernissartensis, I. atherfieldensis.

Sauropoda indet. (? Brachiosauridae)
Aptian:

Ornithopoda: Iguanodon cf. bernissartensis.
Albian:

Sables verts:

Theropoda: Erectopus superbus, Theropoda indet.

Sauropoda indet.

Ankylosauria: Nodosauridae indet.

Pterosauria: Ornithocheiridae indet., Azhdarchidae indet.

Gault Clay, Argiles tégulines:
Sauropoda indet.
Pterosauria: Ornithocheiridae indet.

The available material often (but not always) consists of isolated elements. Apart from some of the
Barremian specimens, most of the Early Cretaceous dinosaur finds from the eastern Paris Basin can be
considered as remains of carcasses derived from nearby land masses (French Central Massif, London-Brabant
Massif) and ultimately buried in marine sediments. The number of taxa represented is relatively modest, but
some trends in the evolution of faunal assemblages may be recognisable. In particular, iguanodontid ornithopods,
which are present in the Hauterivian, Barremian and Aptian formations, have not been reported from the Albian
deposits, despite the fact the latter have yielded a fairly large number of dinosaur specimens. Conversely,
sauropods seem to be relatively more abundant in Albian deposits than in the older formations. The presence in
the Albian Sables Verts of one of the earliest known representatives of the pterosaur family Azhdarchidae is
worth mentioning.

Key words: Early Cretaceous, Paris Basin, Dinosauria, Pterosauria

21

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE HISTORICAL SIGNIFICANCE OF THE IGUANODON
DISCOVERIES AT BERNISSART

Eric BUFFETAUT

Centre National de la Recherche Scientifique (UMR 8538), Laboratoire de Géologie de l'Ecole Normale
Supérieure, 24 rue Lhomond, 75231 Paris Cedex 05, France, eric.buffetaut@wanadoo.fr

The ornithopod dinosaur Iguanodon was first described by Gideon Mantell on the basis of fragmentary
material from the Wealden of Tilgate Forest (Sussex, southern England) in 1824. Although subsequent finds,
notably that of the partial skeleton from Maidstone, contributed to a better understanding of Iguanodon, its
osteology long remained very incompletely known, and its general appearance remained poorly understood, as
shown by the iguana-like reconstructions originally proposed by Mantell and the later reconstructions as a
heavily-built, horned quadruped put forward under Richard Owen's supervision in the 1850s.
The discovery of complete Iguanodon skeletons in the Bernissart coal mine in 1878 led to a thorough
reappraisal of the anatomy of this dinosaur, thanks mainly to the work of Louis Dollo. It became possible to
study many hitherto unknown aspects of the osteology of Iguanodon. This led to Dollo's well-known
reconstruction of a bipedal Iguanodon, with a kangaroo-like stance, as exemplified by the mounted skeletons at
the Brussels Museum and by countless illustrations in papers and books. Although Dollo's conceptions have
been shown to be incorrect in various respects, his palaeobiological approach was in many ways innovative, and
they definitely marked an advance over the earlier views based on incomplete material from England. This new
image of Iguanodon contributed greatly to a better, more realistic, interpretation of dinosaurs as a whole, which
was complemented during the last decades of the 19th century by the finds of well preserved skeletons of various
Jurassic and Cretaceous dinosaurs in western North America. The Bernissart discoveries can thus be considered
as a turning point in our knowledge of dinosaurs.
It should not be overlooked that the excavations in the Bernissart mine brought to light not only the
remarkable collection of Iguanodon skeletons now displayed in Brussels, but also a wealth of other fossils,
including often well preserved remains of plants, insects, fishes, amphibians, turtles and crocodiles. This made it
possible to attempt reconstructions of the Wealden environment in which Iguanodon lived. Although the
conditions under which the fossil accumulation at Bernissart was formed have been the subject of much
controversy, the study of this exceptional locality has contributed significantly to a better understanding of
Wealden environments.
Jules Créteur's discovery of the first Iguanodon bones in the Fosse Sainte-Barbe in March 1878 thus
clearly was the beginning of a major episode in the history of vertebrate palaeontology.

Key words: Bernissart, Iguanodon, History of Palaeontology

22

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
EARLY CRETACEOUS FLORAS FROM SIBERIA, MONGOLIA,
AND NORTH-EASTERN CHINA

Eugenia V. BUGDAEVA, Valentina S. MARKEVICH

Institute of Biology and Soil Science, FEBRAS, 159, Prosp. 100-letiya, Vladivostok, 690022,
bugdaeva@ibss.dvo.ru

Volcanogenic-sedimentary deposits containing abundant remains of fresh-water invertebrates,
vertebrates and floras are widely distributed on the territory of Transbaikalia (Russia), Mongolia and North-
Eastern China. Such fossil assemblages were first discovered during the 19th century by A.P. Gerassimov in the
so called "fish shales" of Turga and Vitim Basins in Transbaikalia. He passed these findings to Academician
A.F. Middendorf and fishes (Lycoptera middendorfii), insects (Ephemeropsis trisetalis), ostracods,
conchostracans (Bairdestheria middendorfii), and mollusks from these Russian localities were subsequently
described. Similar fossil localities were later discovered in Mongolia and North-Eastern China. In China, thesh
typical fossil assemblages, with a high level of endemism, were named `Jehol (Rehe) biota' by alaeontologists.
The attention of the palaeontological society, but also of the public was attracted at the end of the 20th Century,
when feathered dinosaurs, early birds, and ancient angiosperms were unearthed in the Yixian Formation of West
Liaoning Province. The Yixian Formation is unique in the world because of the abundance, biodiversity and
exceptional preservation of the recovered fossils. However, its geological age is still debated.
Jurassic to Aptian ages have been assigned to this stratigraphical unit. According to Chinese
stratigraphers, the age of sediments of Yixian Formation is Late Jurassic (Chen P.-J., Chang Zh.-L., 1994; Cao
Zh.-Y. et al., 1998; Lo Ch.-H. et al., 1999; Sun Ge et al., 1998; Sun Ge et al., 2001), although some
palaeontologists consider it as Early Cretaceous (Li W.-B., Liu Z.-S., 1994,1999; Mao Zh.-Zh. et al., 1990; Pu
R.-G., Wu H.-Zh., 1992; Swisher et al., 1999).
40Ar/39Ar analysis of volcanic rocks frm the Yixian Fm showed values from 121,1±0,2 to 122,9±0,3
million years (Smith et al., 1995). The authors of this paper regard these beds as Barremian in age, although this
interval corresponds in fact to the Aptian (Gradstein et al., 2004).
The fossil flora of the Yixian Formation has much in common with the Barremian-Aptian flora of
Transbaikalia and Mongolia. The common plants are Botrychites reheensis Wu, Neozamites verchojanensis
Vachr., Baikalophyllum lobatum Bugd., Pityolepis pseudotsugaoides Sun et Zheng, Podocarpidites reheensis
(Wu), Brachyphyllum longispicum Sun, Zheng et Mei, Scarburgia hilii Harris, Baikalophyllum lobatum Bugd.,
Baiera valida Sun et Zheng, cf. Lindleycladus lanceolatus (L. et H.) Harris, Ephedrites chenii (Cao et Wu),
Carpolithus multiseminalis Sun et Zheng, C. pachythelis Sun et Zheng, Baisia hirsuta Krassil.
The Barremian-Aptian flora of Transbaikalia is highly diversified, cycadophytes and various conifers
being particularly well represented. Principal index fossils are Baisia hirsuta, Pseudolarix and the bennettite
Otozamites lacustris.
The high diversity of this flora facilitates correlation with neighboring regions. Related floras occur in
the Barremian-Aptian Chegdomyn and Chemchukin Formations of the Bureya Basin (Far Eastern Russia). The
Ussuri (Barremian) and Lipovtsy (Aptian) Formations of Razdolnoe basin and the Severosuchan Formation
(Aptian) of Partizansk basin in Southern Primorye contain Cladophlebidium dahuricum Pryn., which is typical
for the Transbaikalia flora.
The Transbaikalia flora can also be correlated with the Aptian flora of Yakutia (Siberia). Both floras
contain typical taxa such as Gleichenia lobata Vachr. and Neozamites verchojanensis Vachr.
Four characteristic assemblages are recognized by V.A. Krassilov in the Early Cretaceous of Mongolia:
1) Baiera manchurica (Berriasian?); 2) Otozamites lacustris-Paseudolarix erensis (Barremian-Aptian); 3)
Baierella hastata-Araucaria mongolica (Aptian); 4) Limnothetis-Limnoniobe (Aptian to early Albian(?). Leaves
of Otozamites and Pseudolarix have been found in Transbaikalia flora, and morphological and epidermal studies
show similarities with Otozamites lacustris from Mongolia. These plants permit correlation of Transbaikalia
flora with the Early Cretaceous Mongolian floras from Bon-Tsagan, Manlaj, and Gurvan-Eren. The Yixian and
Mongolian floras also contain common taxa, like Gurvanella, some conifers and gnetophytes.
Therefore, the age of the Yixian Formation is not, in our opinion Late Jurassic or situated at the
boundary between the Jurassic and the Cretaceous. Judging from the palaeobotanical data, we can conclude that
the age of this famous stratigraphical unit is Barremian to Aptian.

Key words: Jehol biota, Transbaikalia, fossil flora

23

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
TO BE OR NOT TO BE: CAN CONODONTS BE VERTEBRATES?

Pierre BULTYNCK

Department of Paleontology Royal Belgian Institute of Natural Sciences, Vautier street 29 B-1000 Brussels
Belgium, pierre.bultynck@naturalsciences.be

Conodonts were originally described by Pander in 1856 as isolated, microscopic fish teeth. Until the
1920ies they did not receive much attention from paleontologists. Already in that first period, conflicting ideas
arose about the taxonomic treatment of conodonts and their zoological affinities. However, these problems were
only thoroughly debated 50 to 80 years later, after conodonts became well known as one of the most important
tools for marine Paleozoic and Triassic biostratigraphy. In 1969, different sets with biostratigraphic subdivisions,
principally from Europe and North America, were presented at a symposium on Conodont Biostratigraphy in
Columbus (Ohio). Two years later the taxonomic problem of artificial form taxonomy (in which definition of
conodont species is based on characteristics of individual conodont elements) versus multi-element taxonomy
was taken up again. The latter is based on findings of natural conodont assemblages on bedding planes. They
show that different form species constitute a bilaterally symmetrical apparatus in the "conodont animal". This
taxonomic problem was discussed at the 1971 Marburg Conodont Symposium and the use of multi-element
taxonomy was recommended. It should be stressed here that reconstruction of conodont apparatuses is not only
important for taxonomy but also for the understanding of the function of this apparatus in the conodont animal.
As such, it is one of the elements to find out the zoological identity of conodonts.
The fortuitous discovery of a conodont animal in the collections of the British Geological Survey in
Edinburgh (+/-1980) marked a new period of intense discussions on the zoological affinities of conodonts. The
4.5 cm long elongated specimen shows preservation of the body outline, some internal soft parts and a natural
assemblage of conodont elements. The specimen originates from the Lower Carboniferous Granton Shrimp bed
near Edinburgh, in which later nine additional specimens were found. The specimens were intensively studied by
British research teams mainly including conodont experts. As the preservation is moderate, interpretation of
structures remains open to discussion. The study of the Granton specimens was combined with that of an Upper
Ordovician species from South Africa, with investigations on the structure of conodont elements and their
histology-histochemistry and the function of the conodont apparatus. Donoghue, Forey & Aldridge (2000)
finally concluded on the basis of a cladistic phylogenetic analysis that conodonts are stem gnathostomes
(vertebrates). However this view was in contradiction with the results of earlier histological-histochemical
analyses of conodont hard tissues. These results did not support vertebrate affinities but indicate protochordate
affinities (Kemp & Nicoll, 1995, 1996; Schulze, 1996). Turner, Blieck & Nowlan (2004) stated that on the basis
of their hard tissues conodonts cannot be considered as vertebrates and they assigned them to "unresolved or
questionable basal chordates". They fully explained their point of view in a manuscript now submitted for
publication. (Turner, Burrow, Schultze, Blieck, Reif, Rexroad & Nowlan False teeth! Why conodonts are not
vertebrates),
Szaniawsky & Bengston (1993) demonstrated the evolution from coniform paraconodonts to coniform
euconodonts (= conodonts as discussed above). Earlier Szaniawski (1987) considered that proto- , para- and
euconodonts are a monophyletic clade that may show affinities with chaetognaths. Several vertebrate experts
have stressed that conodont elements differ from vertebrate teeth by their ultrastructure and chemical
composition. Moreover, I consider that growth and detailed morphology of conodont elements, the architecture
of the conodont apparatus, and the early evolution of that apparatus do no t support a close relationship with
vertebrate teeth. As a closing comment it can be stated that conodonts do not fit well in the early evolutionary
stage of craniates and vertebrates. Some characters are in agreement with a basal chordate position.

Key words: conodonts, taxonomy, function, zoological affinities

24

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE BRAINCASE OF RHABDODON (DINOSAURIA:
ORNITHOPODA): NEW SPECIMENS FROM THE LATE
CRETACEOUS OF SOUTHERN FRANCE

Phornphen CHANTHASIT

UMR 5125 - Paléoenvironnement & Paléobiosphère, Université Claude Bernard Lyon1,
2 rue Raphaël Dubois 69622 CEDEX Villeurbanne, France,
phornphen.chanthasit@pepsmail.univ-lyon1.fr

Rhabdodon priscus was first described from the early Maastrichtian of La Nerthe (Bouches-du-Rhône) by
Matheron in 1896. Rhabdodon fossils have been continued in several Campanian-Maastrichtian localities in
Europe especially in southern France. However, cranial skeletons of Rhabdodon have rarely been found and they
are not well preserved. Particularly the braincase, which provides important morphological characters for
phylogenetic analyses was hardly mentioned. Recently, two new braincases referred to Rhabdodon from a
locality near Cruzy (Hérault) and from a locality in Vitrolles (Bouches-du-Rhône) have been found. They are
more complete and show more braincase elements than the braincases reported in preceding works. These new
braincases are described herein and compared to those of Zalmoxes (another Rhabdodontidae) and those of the
other euornithopod dinosaurs described in the literature.

Key words: dinosauria, ornithopoda, Rhabdodon, braincase, France

25

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
NECK UTILIZATION AND FEEDING STRATEGY IN SAUROPOD
DINOSAURS

Andreas CHRISTIAN, Gordon DZEMSKI, Jan-Thomas MÖLLER

1 Universität Flensburg, Institut für Biologie und Sachunterricht und ihre Didaktik, Auf dem Campus 1,
24943 Flensburg, Germany, christian@uni-flensburg.de

Based on a comparison of stress values in the intervertebral discs along the vertebral column of the neck,
the neck posture is reconstructed for different sauropods. For each specimen, the range of possible stress values
in the intervertebral discs is tested for different neck postures in order to find habitual postures which are
characterised by more or less constant stress values along the neck. In order to reconstruct neck movements,
lever arms of muscles, tendons, and ligaments and the articulation of adjacent neck vertebrae are analysed in
some specimens. For comparison, neck anatomy and neck movements are studied in long-necked vertebrates like
giraffes, camels and ostriches. For sauropods of different sizes, the energetic costs of neck movements are
estimated and compared with the energy expenditures of locomotion and the basal metabolism. The results
indicate considerable differences in the posture and utilization of the neck among sauropods. Some sauropods,
like Brachiosaurus and Euhelopus, used the neck in a similar way as giraffes do, whereas the feeding strategy of
other species, like Diplodocus and Apatosaurus, was more like in camels or ostriches. In a single sauropod, the
neck posture may have varied considerably during different activities like standing at rest, locomotion or
feeding. Vertical changes of the head during feeding depended on the spatial distribution of the food. For very
large sauropods extensive vertical movements of the neck or of the whole body during feeding were efficient
only if the sources of food were spaced widely apart.

Key words: sauropods, neck, feeding

26

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
WHAT'S NEW IN THE WORLD OF DEVONIAN TETRAPODS?

Jennifer A. CLACK

University Museum of Zoology, Cambridge, Downing St., Cambridge CB2 3EJ UK, j.a.clack@zoo.cam.ac.uk

The past 15 years has seen an exponential increase in the number of known Devonian tetrapod taxa, with
11 genera now named and other material awaiting description. Since 1993 Devonian tetrapods have been
discovered in North America, Scotland, Latvia, China and Belgium, and further finds from other Russian
localities have also been reported. Correspondingly there has been an expansion in the geographical and
environmental range that they are known to have inhabited. Thus by the end of the Devonian they are known to
have occurred in most of the major continental masses, including Gondwana. The environments that the tetrapods
inhabited range from fully freshwater, to marginal marine, encompassing estuarine and lagoonal sites. The
implication here is that the earliest tetrapods were euryhaline and could move around continents along their
margins or through shallow interconnecting seas.

The mid- to Late-Devonian was period of great environmental change: oxgyen levels dropped to an
estimated 13%; temperatures and carbon dioxide levels were both elevated relative to today; and it was during this
period that terrestrial plants increased in size, diversity and range. For the first time, decaying organic matter in
the water contributed to widespread anoxia in both marine and riverine systems. Yet this was the time during
which the tetrapod stem group also diversified, and it is notable that modifications to breathing and support
apparatuses were among the first to occur. Recent finds of Late Devonian tetrapod-like fish and tetrapods with
limbs are serving to close the morphological gap between finned and limbed tetrapodomorphs and are beginning
to show the sequence of acquistion of such characters in the origin of limbed tetrapods. The picture is very
different from scenarios generated in the early 20th century when information was scarce.

Among the most recent developments is the description of Ventastega as one of the most primitive
tetrapods: in having an enlarged spiracular notch, it resembles the tetrapod-like fish Tiktaalik from the Frasnian of
Canada. Its limb girdles are similar to those of Acanthostega implying the presence of limbs with digits, but it has
some more primitive features in its lower jaw and braincase. Its occurrence in Laurussia is consistent with a
Laurussian origin for limbed tetrapods.

The iconic genus Ichthyostega has been restudied, showing that its ear region was probably adapted for
underwater audition, whereas its postcranial skeleton shows a mixture of aquatic (paddle-like hind limb, finned
tail) with terrestrial (robust shoulder and forelimb, differentiated axial skeleton with broad thoracic ribs)
adaptations. Work is planned to investigate this by creating a virtual computer model of its skeleton. Subadult
humeri of Ichthyostega appear to retain more primitive fish-like features than the equivalent stage in
Acanthostega: this has locomotory, ontogenetic, and phylogenetic implications. Discovery of an ichthyostegid
outside East Greenland, in Belgium, has biogeographical significance.

Much of the recent information has been gathered by the use of computer-assisted tomography (CT
scanning), which is allowing unprecedented access to previously intractable specimens. Application of such
techiques at finer and finer resolutions will allow us to pose and answer questions that would have been
considered impossible by earlier workers, such as the position, size, and direction of muscle attachments. All of
this will help to build an even more complete picture of the transition from fish to tetrapod and from water to land.

Key words: Devonian, tetrapods, evolution

27

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
BIODIVERSITY OF THE DEVONIAN TETRAPOD-BEARING
LOCALITY OF STRUD, BELGIUM

Gaël CLEMENT 1, Cyrille PRESTIANNI 2

1 Département Histoire de la Terre, UMR 5143 du CNRS, Muséum national d'Histoire naturelle, 57 rue
Cuvier, 75231 Paris cedex 05, France, gclement@mnhn.fr
2 Unité de Paléobotanique - Paléopalynologie - Micropaléontologie (PPM), Département de Géologie,
Université de Liège, Allée du 6 Août, Bâtiment B-18, 4000 Liège 1, Belgium, cyrille.prestianni@ulg.ac.be

The origin of tetrapods (four-legged vertebrates), about 365 million years ago, was one of the key
events in the evolution of vertebrates. It changed the terrestrial ecosystem forever and gave rise to a major new
group of animals that today numbers some 24,000 living species. In contrast to recent morphological and
phylogenetic advances, there remains a serious gap in our knowledge of the evolutionary, ecological and
biogeographical context for the origin of tetrapods. Few Devonian tetrapod-bearing localities allow exhaustive
environmental and ecological studies. The Strud quarry (Namur Province, Belgium) was first investigated by
Hock (1878) and plant and vertebrate fossil material from Strud was later published by Lohest (1888), Leriche
(1931) and Stockmans (1948). Unfortunately the precise location of the site was lost for more than a century. An
incomplete lower jaw, first described as a fish remain by Lohest (1888), has been recently determined as a
tetrapod mandible. Following this discovery, the Strud locality was rediscovered (2005) and has yielded a
diverse Famennian (Late Devonian) flora and fauna. It includes miospores, plant megafossils, arthropods and
vertebrates. The new material is abundant and highly valuable. Although mainly disarticulated, fossil remains
from Strud are regarded as autochthonous. The quality of the material allows accurate identifications. Bone is
well-preserved and delicate elements such as Phyllolepis plates or dentary teeth on lower jaws are always found
unbroken. The CTscan technique has been used to perform virtual 3D reconstructions of remains enclosed in the
fine-grained sandstone (e.g., lungfish and tetrapod lower jaws). The exquisite preservation of arthropods and
plants in the highest layers of the section points to a reduced post-mortem transport, allowing to consider the
taphonomic assemblage as a biological community. Amongst others, key taxa of the Strud locality are: earliest
demonstrated pre-ovule plants Moresnetia, Pseudosporogonites, and Condrusia, exquisitely preserved
malacostracan and Triops-like notostracan crustaceans, rhynchodipterid lungfishes, rhizodontid tetrapodomorph
fishes and Ichthyostega-like tetrapods.
Preliminary results indicate that the miospore assemblage is characteristic of the GF (Grandispora
gracilis - Grandispora famennensis) biozone (late mid Famennian). This indicates that the Strud locality is
slightly older than the Famennian tetrapod sites of East Greenland (Late Famennian), and is the oldest locality
yielding Moresnetia.
The depositional environment of the Late Devonian Strud locality, preliminary defined as a flood facies
on an alluvial plain, is rich in crucial data (traces of emersion, of cyclicity, of warm climate, etc.). These deposits
are compared with those of the outstanding Famennian tetrapod-bearing localities of East Greenland (inland
fluviatile floodplain facies) and Pennsylvania (estuarine floodplain facies). The vertebrate faunal assemblage
comparisons of these three localities show considerable faunal overlap at genus level although some remarkable
differences also occur. If the Strud locality is confirmed as Late Middle Famennian in age, Belgian occurrences
of Gondwanan taxa are then the very first ones in Euramerica. It is for instance the case for the placoderm
Groenlandaspis, the lungfish Soederberghia, and the rhizodontid tetrapodomorphs, restricted to marginal or
freshwater environments and known to have been globally distributed during the Late Famennian times. These
records seem to contradict the previous hypothesis of a wide separation between Euramerica and Gondwana. A
strange fact, and possibly important, is the strong imbalance between the high number of vertebrate gondwanan
taxa in Euramerica at the end of the Famennian compared to the low number of Euramerican taxa in Gondwana.
This interpretation has been recently called the "Great Devonian Interchange" and is still poorly understood.

Key words: Belgium, Late Devonian, Paleobiogeography, Paleoenvironment, Seed plants, Vertebrates

28

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
MAASTRICHTIAN LAND VERTEBRATES IN RUSCA MONTAN
BASIN (ROMANIA)

Vlad CODREA 1, Pascal GODEFROIT 2, Thierry SMITH 2, Ctlin JIPA-MURZEA 1

1 Babe -Bolyai University, Department of Geology-Paleontology, 1 Koglniceanu Str., 400084 Cluj-Napoca,
Romania, codrea_vlad@yahoo.fr ; vcodrea@bioge.ubbcluj.ro
2 Institut Royal des Sciences Naturelles de Belgique, Department de Paléontologie, Rue Vautier 29, B-1000
Bruxelles, Pascal.Godefroit@naturalsciences.be ; Thierry.Smith@naturalsciences.be

The Rusca Montana Basin is located in Poiana Rusc Mountains (Occidental Carpathians, Romania),
south of Mure River. The basin has a triangle-shaped elongated outline, trended on a westeastern direction. Its
basement involves metamorphic and Upper Cretaceous (Santonian-Early Maastrichtian) flysh deposits. The thick
(around 2500 m) basin filling sediments pile (Late Cretaceous-? Paleogene) forms an asymmetrical syncline,
framed to the north by an important fault. The basin formed after the Late Cretaceous "Laramian" tectogenesis
(Sndulescu, 1984), in rather similar way as the neighboring Haeg Basin (Willingshofer et al., 2001). Like in
Haeg Basin, the infilling deposits started to accumulate in the Latest Cretaceous (Maastrichtian) and form thick
continental sequences. However, there are important geological differences between these basins: Upper
Maastricthian coal deposits were mined in Rusca Montan for several decades in the last century and geological
history of this basin comprises magmatic events related to banatites (volcanic breccia, agglomerates, lava flows,
volcanic tuffs, dykes and sills maturing the coal, where in contact). In Haeg Basin, the magmatism inference is
weaker, and mainly concerns the Densu -Ciula Formation.
The first geological studies in Rusca Montan begun in 19th century, but its real interest increased only
when the coal started to be mined in the third decade of the 20th century. The coal-bearing strata yielded rich
macroflora assemblages, including several Pandanus species, ferns, angiosperms (Schafarzik, 1907; Tuzson,
1913; Givulescu 1966, 1968; Petrescu & Du a, 1984), but also microflorae (Antonescu et al., 1983).
Although some geologists suggested the existence of coeval vertebrates too (e.g. Mamulea, 1952; Dinc
et al., 1971; Dinc, 1977), vertebrate fossil have not been reported up to now in this area. Even the notorious
Nopcsa's (1905) geological map does not report any vertebrate find in this basin. In the last decade, on the
easternmost side of the basin (Lunca Cernii), a small outcrop exposing a short sequence of Maastrichtian
continental deposits of fluvial origin (red siltic clay, coarse sand and microconglomerate channel fills) yielded
some teeth and bones belonging to dinosaurs (sauropod titanosaur, the euornithopod Zalmoxes, Velociraptorinae
indet., Troodontidae indet., other Theropoda indet), turtles (? Kallokibotion), indeterminate crocodilians and
multituberculate mammals (Kogaionidae). These fossils are reported for the first time in this communication.
Besides vertebrates, these deposits have also yielded abundant vegetal remains (mainly seeds), small red-
yellowish amber pieces, and rare invertebrates (snails, ostracods...).
Both the lithology and the fauna are closely similar to those from the Sânpetru and Densu -Ciula formations in
Haeg Basin. One can presume that Rusca Montan Basin is nothing else but a western extension of a larger
Maastricthian sedimentary area, which once included the Haeg Basin too. Both basins were depositional centers
for the detrital rocks resulted from the erosion of the "post-Laramian" sedimentary cover spread on large areas,
formed on the new thrusted nappes in Occidental Carpathians. These detrital rocks had been reworked by the
fluvial systems and preserved in these basins where erosion did not act in an aggressive way. If these two basins
are now separed, it is due to the subsequent uplifts of the metamorphic tectonic blocks in Poiana Rusca
Mountains during the Cenozoic.

29

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
SPECIALIZED DENTITION IN LOWER CRETACEOUS
FRESHWATER HYBODONT SHARKS

Gilles CUNY 1, Lionel CAVIN 2, Varavudh SUTEETHORN 3

1 Natural History Museum of Denmark, University of Copenhagen, Øster Voldgade 5-7, 1350 Copenhagen K,
Denmark, gilles@snm.ku.dk
2 Department of Geology and Palaeontology, Muséum d'Histoire naturelle, CP 6434, 1211 Genève 6,
Switzerland, lionel.cavin@ville-ge.ch
3 Department of Mineral Resources, Rama VI Road, Bangkok 10400, Thailand, suteethorn@hotmail.com

A new species of hybodont shark with a cutting dentition, Mukdahanodus trisivakulii gen. nov. et sp.
nov., from the Lower Cretaceous Sao Khua Formation of Thailand has recently been described. After a turn-over
in hybodont faunas in Thailand, it was apparently replaced in the Aptian/Albian ecosystem by Thaiodus ruchae,
which independently developed a very similar dentition. Thaiodus and Mukdahanodus belong however to two
separate families. A comparative study indicates that cutting dentitions with serrated teeth appeared four times
independently within the hybodont sharks over a rather short period of time, from the Late Jurassic to the Albian.
Moreover, such a dentition occurred only in species spending at least part of their life cycle in fresh waters.
Hybodont sharks with a cutting dentition seem to have independently evolved in Southeast Asia
(Mukdahanodusand Thaiodus) and in the Africa-South America continent (Pororhiza and Priohybodus). Two
main kinds of cutting dentition can be identified among hybodonts: high-crowned (Priohybodus) and low-
crowned (Mukdahanodus gen. nov., Thaiodus and Pororhiza). The development of these cutting dentitions in
hybodont sharks is linked to a compaction of their enameloid microstructure.

Key words: Hybodont sharks, Cretaceous, cutting dentition

30

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
PALYNOLOGICAL OVERVIEW OF WEALDEN FACIES SEDIMENTS
FROM BELGIUM AND NORTHERN FRANCE

Jean DEJAX 1, Denise PONS 2, Johan YANS 3

1 MNHN, Département "Histoire de la Terre", Paris, France, dejax@mnhn.fr
2 Université Pierre et Marie Curie-Paris 6, Paris, France, Denise.pons@upmc.fr
3 FUNDP Geology, UCL-Namur, Belgium, Johan.Yans@fundp.ac.be

Several palynological studies have dealt with Wealden facies sediments from the northern part of
Europe since the pioneering work of Delcourt & Sprumont (1955). Beginning with Bernissart famous buried
locality (Yans et al., 2005; Dejax et al., 2007a), we investigated clays from Baudour and Hautrage quarries
(Dejax et al., 2007b, 2008). We now present some new results from drillings at Bernissart (BER-3) and Baudour.
Leaving the Mons Basin, we also present results from Soignies karst filling (in the Namur Synclinorium also
called "Parautochtone Brabançon") and from the Longueville quarry (Pas-de-Calais, northern France) which was
also a pioneering study (Herngreen, 1971). Each locality yielded a rich microflora entirely continental in origin.
Fern spores and bisaccate pollen grains are a main feature of the assemblages, allowing deductions about the
climate and the environment; less numerous, angiospermous pollen grains are also present, mostly known as
biorecords sensu Hughes' peculiar terminology (Hughes, 1976, 1994; Hughes et al., 1979). These biorecords are
important, as palynological milestones of angiosperm evolution as well as stratigraphic guide forms which allow
correlations within the English Weald and Wessex Basins, where the continental Lower Cretaceous is divided
into six MCT (Monosulcate Columellate Tectate) phases, ranging from Hauterivian to Aptian.

DEJAX J., PONS D., YANS J. (2007a). Palynology of the dinosaur-bearing Wealden facies in the natural pit of Bernissart (Belgium). Rev.
Palaeobot. Palynol., 144, 25-38.
DEJAX J., DUMAX E., DAMBLON F., YANS J. (2007b). Palynology of Baudour Clays Formation (Mons Basin, Belgium): correlation
within the "stratotypic" Wealden. In: Steemans P., Javaux E. (eds.), Recent Advances in Palynology. Carnets de Géologie /
Notebooks on Geology, Memoir 2007/01, 16-28.
DEJAX J., PONS D., YANS J. (2008). Palynology of the Wealden facies from Hautrage quarry (Mons Basin, Belgium). Mem. Geol. Survey
of Belgium, 55, 45-51.
DELCOURT A., Sprumont G. (1955). Les spores et grains de pollen du Wealdien du Hainaut. Mém. Soc. belge Géol., Paléont. Hydr., 5, 1-
73.
HERNGREEN G.F.W. (1971). Palynology of a Wealden section (Lower Cretaceous) in the "Carrière de Longueville", the Boulonnais
(France). Rev. Palaeobot. Palynol., 12, 271-302.
HUGHES N.F. (1976). Palaeobiology of angiosperm origins. Cambridge University Press, 242 p.
HUGHES N.F. (1994). The enigma of angiosperm origins. Cambridge University Press, 303 p.
HUGHES N.F., DREWRY G.E., LAING J.F. (1979). Barremian earliest angiosperm pollen. Palaeontology, 22/3, 513-535.
YANS Y., DEJAX J., PONS D., DUPUIS C., TAQUET, P. (2005). Implications paléontologiques et géodynamiques de la datation
palynologique des sédiments à faciès wealdien de Bernissart (bassin de Mons, Belgique). C. R. Palevol 4, 135-150.

Key words: Bernissart, Baudour, Soignies, Longueville, Wealden facies, palynology

31

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
FROM LAND TO SEA: THE EARLY EVOLUTION OF CETACEANS

Christian DE MUIZON 1, Jonathan GEISLER 2

1 UMR 7207, Département Histoire de la Terre (CP 38), Muséum National d'Histoire Naturelle, 57, rue
Cuvier, PARIS F-75005, France, muizon@mnhn.fr
2 Department of Geology and Geography, Georgia Southern University, Statesboro, Georgia 30460-8149,
USA, geislerj@georgiasouthern.edu

Cetaceans are the most highly modified mammals and their origin has always been enigmatic in the last
centuries. It is now unanimously accepted that they originate among terrestrial artiodactyls during the early
Eocene. Molecular studies regard the family Hippopotamidae as the sister group of Cetaceans. Morphological
analyses (therefore, including fossils) are not consensual. Some confirm the molecular studies, others regard the
Hippopotamidae as the sister group of the exclusively fossil anthracotheres, this monophyletic group
representing in turn the sister group of cetaceans. Affinities of cetaceans have recently been unequivocally
confirmed by discoveries of spectacular fossil in Pakistan (Pakicetus and Rodhocetus), which bear the
characteristic double-pulleyed astragalus of artiodactyls. Therefore, the fossil ungulates, Mesonychians, which
have a simple-pulleyed astragalus, cannot be the sister group of cetaceans anymore as they had been regarded for
several decades.
The oldest known cetacean is Pakicetus, a semi-aquatic cetartiodactyl from the early Eocene of
Pakistan. Pakicetus was largely terrestrial (possibly cursorial), but entered the water in search for food or
possibly to protect its skin from the sun or to prevent any other kind of danger. Pakicetus had osteosclerotic
bones, which are characteristic of semi-aquatic mammals. At the same epoch in Pakistan, Indohyus was a semi-
aquatic artiodactyl of the family Raoellidae, which is regarded as the sister group of cetacean, but which may
also be included in the cetaceans as well. A few million years later, Ambulocetus was an amphibious cetacean
capable of moving on land but also an agile swimmer using its hind limbs for propulsion. Ambulocetus was a
formidable predator with powerful teeth. During the middle Eocene protocetids are other partly amphibious
cetaceans, although some of them were probably hardly capable to move on land. Rodhocetus, remingtonocetids,
Georgiacetus, still had well-developed limbs but they were progressively loosing their links with land. Their
nares significantly migrated backward (those of Pakicetus and Ambulocetus were more or less apical) and their
inner ear acquired adaptation to aquatic life.
During the late Eocene, appeared the first strictly aquatic and highly pelagic cetaceans, the
Basilosauridae. Their hind limbs are totally atrophied and are not functional. The nares are on the dorsal face of
the rostrum, on the anterior third of the skull. From that time the way toward modern cetaceans was open: the
earliest mysticetes are from the latest Eocene of Antarctica and the oldest odontocetes are from the early
Oligocene.

Key words: Cetaceans, Eocene, evolution, aquatic life

32

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
VERTEBRATE REMAINS IN THE 2003 BERNISSART DRILL:
HISTOLOGICAL ASSESSMENT

Armand DE RICQLES 1, Pascal GODEFROIT 2, Johan YANS 3

1 Collège de France, / (UMR 7072 CNRS-UPMC-UCP)
UPMC Paris Universitas, France, armand.de_ricqlès@upmc.fr
2 Department of palaeontology, Royal Belgian Institute of Natural Science, Rue Vautier, 29
1000 Bruxelles, Belgium , Pascal.Godefroit@naturalsciences.be
3 Facultés Universitaires de Namur, Belgium, jyans@fundp.ac.be

Following the 2003 drill in Bernissart (BER 3 borehole), examination of the column revealed stony dark
greyish remains at levels -296,5m and -309m respectively, hence within the Wealden clays levels.
Four fragments of a few CC each were carefully extracted for further examination by histological
technics. The fragments, numbered Bernissart 3 296,5, Bernissart 3 309 A, Bernissart 3 309 B, Bernissart 3
309 C were dried and resin embedded under gentle vacuum, the resin temperature being monitored to secure a
slow polymerization. The resulting blocks were trimed and sawed on a thin diamond/copper circular blade and
further processed to obtain thin sections following the routine palaeohistological technics.
For comparative purposes, some bone material (rib, specimen H) as well as a maxillary tooth from a
Bernissart Iguanodon were obtained from the RBINS collection. Maxillary and dentary teeth from Bactrosaurus
were also histologically processed. The resulting 26 thin sections were examined under the dissecting and the
compound microscope, in ordinary and polarized light.
Bernissart 3 296, 5 specimen. All the sections show compact bone fragments around a small cavity
filled by a black material containing some teeth remains. The bone fragments have a complex structure of
compacted secondary endosteal trabeculae and Haversian systems. Lack of Sharpey's fibers preclude that the
bone tissues are actually periosteal or dermal, although the bones they come from may well have had such an
ontogenetic origin (dentary, maxillary...). The tooth remains suggest thin elongate small teeth with a rather
pointed tip. The enamel is thin and non prismatic. The dentine shows the traditional radially oriented canaliculi.
No root system or ankylosis on dentigerous bone could be observed.
Bernissart 3 309 A, B, C specimens all show a more or less dense Haversian bone tissue intimately
associated with massive pyrite deposition. Some regions apparently preserve the natural free surface of bone and
it is possible to observe there some primary (periosteal) bone tissue, more or less invaded by secondary osteons
(Haversian systems).

Discussion. The bone tissues observed at level 309 agree well with previous descriptions of adult
dinosaurian bone tissues (dense Haversian bone) in general, and notably with Iguanodon. Direct comparison
with a small Iguanodon rib (specimen H, diameter :12x20 mm) even suggest that the 309 m drill material may
have pertained to a "early adult " stage, according to the incomplete Haversian replacement in the superficial
region, occurrence of LAGs in primary bone and poor vascularity of the later, although a clear EFS was not
observed. Lack of radial "craks" at the periphery of the secondary osteons would not suggest an early aquatic
taphonomic episode. The teeth observed at level 296,5 m clearly differ in size and structure from the ones of
Iguanodon and other ornithopods. Their small size, likely slenderness and thin enamel do not fit with crushing
functions. Unfortunately, lack on information on their mode of ankylosis and on the occurrence of a root
precludes further diagnosis. There are also no clear evidences of actual (anatomical) relationships with the
surrounding bone fragments. Whatever it may be, the lucky occurrence of bone and tooth material at two
superposed levels in a small diameter drill underline again the fossil abundance and value of the Bernissart
Wealden pit.

Key words: bone and tooth histology, Iguanodon, dinosauria, other bony vertebrates remains

33

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
IGUANODON-LIKE FOOTPRINTS FROM THE ENCISO GROUP
(APTIAN, LOWER CRETACEOUS) OF LA RIOJA (CAMEROS BASIN,
SPAIN)

Ignacio DÍAZ-MARTÍNEZ 1,2,4, Félix PÉREZ-LORENTE 1,2, Xabier PEREDA-
SUBERBIOLA 3, José Ignacio CANUDO 4

1 Fundación Patrimonio Paleontológico de La Rioja, Portillo 3, 26586 Enciso, Spain, inaportu@hotmail.com
2 Edificio C. T. U. de La Rioja. Madre de Dios, 51. 26006, Logroño, Spain, felix.perez@unirioja.es
3 Universidad del País Vasco/EHU, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y
Paleontología, Apartado 644, 48080, Bilbao, Spain, xabier.pereda@ehu.es
4 Grupo Aragosaurus, Paleontología, Facultad de Ciencias, Universidad de Zaragoza, Pedro Cerbuna 12,
50009 Zaragoza, Spain, jicanudo@unizar.es

Large ornithopod (Iguanodon-like) footprints are relatively abundant in the Enciso Group of the
Cameros Basin (La Rioja, Spain). Casanovas & Santafé (1971 and subsequent works) referred these tracks to
Iguanodon or to "iguanodontid" ornithopods. A revision of the La Rioja footprints allows providing information
about the palaeobiodiversity of the trackmakers.
Moratalla (1993) distinguished three morphotypes of Iguanodon-like footprints, which are present in La
Magdalena (Préjano), Peña Untura (San Román de Cameros) and Valdeté (Préjano) sites. In this work a fourth
morphotype (Hadrosaurichnoides) from La Era del Peladillo site (Igea) is also considered. All of them are
known in the Enciso Group (Aptian).
Morphotype 1. "Brachyiguanodonipus". It represents graviportal, mesaxonic and tridactyl footprints.
Wider than long, with a rhomboidal pad per digit, and other sub-rounded and wide in the heel. Short digits, as
long as wide, with a rounded to blunt distal ending. Notch present in the proximal part of the digits II and IV.
Morphotype 2. "Iguanodonipus". This morphotype grouped mesaxonic and tridactyl footprints with
similar length and width, with rounded and wide hell. Sometimes the posterior part of the heel is narrow and
elongated, probably for the metatarsal support. The digits are longer than wide, sub-triangular, with sub-acute
distal ends. The notch (if present) is not well marked.
Morphotype 3. Indeterminate ichnotaxon. This morphotype grouped mesaxonic and tridactyl footprints.
They are longer than wide, with a pad per digit and other in the heel. The digits are longer than wide, with a
blunt distal ending. This indeterminate taxon present well marked notchs and short and rounded heel.
Morphotype 4. "Hadrosaurichnoides". It represents to mesaxonic and tridactyl footprints with
approximately as long as wide, with a pad per digit and other in the heel. The digits are short and wide, with a
blunt distal ending. "Hadrosaurichnoides" present interdigital web and short and rounded heel, without notch.
The shape differences in the footprints from the Lower Cretaceous Cameros Basin suggest that four
different ichnotaxons are represented. The morphology of the footprints 1, 3 and 4 is coherent with that of
Iguanodon or closely related large iguanodontoids. On the basis of the morphotype 2, the presence of more basal
ornithopods (basal ankylopollexians?) cannot be rejected.

Key words: Iguanodon-like, footprints, La Rioja, Cameros Basin, Spain

34

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
TRANSMISSION ELECTRON MICROSCOPY OF SAUROPOD BONES:
A LOOK AT THE NANOSTRUCTURE

Maïtena DUMONT 1, András BORBELY 1, Aleksander KOSTKA 1, Anke PYZALLA 2

1 Max Planck Institut, Max-Planck Strasse 1 - 40237 Düsseldorf - Germany, m.dumont@mpie.de
2 HelmholtzZentrum Berlin, Glienicker Strasse 100, 14109 Berlin - Germany, anke.pyzalla@helmholtz-
berlin.de

Bone has a hierarchical structure composed of an arrangement of materials at many length scales, which
work in concert to achieve excellent mechanical properties. At the lower nanostructural level bone is composed
of collagen fibrils as a matrix reinforced with mineral particles. The manner in which these two components are
organized has important consequences for the properties of the structure. So far the characterization of
sauropodomorph fossil bones has been concentrated mainly on the microstructural level, previous works
showing that sauropod bones have a similar fibrolamellar bone tissue as mammals (cow, elephant...). In the
present study the nanostructure of the sauropod bone has been analysed by transmission electron microscopy
(TEM) and compared with cow bone. This technique coupled with electron diffraction provides the
determination the dimensions and the shapes of bone apatite nanocrystals. Both sauropod and cow samples for
TEM investigations were prepared with a Focus Ion beam (FIB) system.
The aim of this work is to enlighten the potential of TEM in providing precise, localized size analysis
but also information concerning the effect of diagenesis in fossil bone at the nanolevel.

Key words: TEM, bone structure, sauropod, nanocrystals

35

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
PTEROSAUR FLIGHT DYNAMICS AND THE EVOLUTION OF
FLIGHT STYLE WITHIN THE LINEAGE: INITIAL EXPERIMENTAL
FINDINGS

Ross ELGIN 1, Edina PRONDVAI 1, Eberhard FREY 1, Wolf KRÜGER 2

1 Staatliches Museum für Naturkunde Karlsruhe (SMNK), Erbprinzenstraße 13, D-76133 Karlsruhe,
rosselgin@hotmail.com
2 Deutsches Zentrum für Luft- und Raumfahrt (DLR), Institut für Aeroelastik, Bunsenstr. 10, D-37073
Göttingen, wolf.krueger@dlr.de

The pterosaurs were a group of Mesozoic reptiles that originated from a still uncertain ancestor in the
Triassic and persisted until the very end of the Cretaceous. They represented an important step in vertebrate
evolution, being the first vertebrates to evolve true powered flight, while some Late Cretaceous forms were
capable of growing to gigantic sizes (~10m wingspan) making them the largest animals to ever take to the air. A
hypothetical evolutionary pathway can be reconstructed from a simple arboreal glider to the wide variety of
flight styles (eg. marine/dynamic soaring, thermal soaring) that must have existed within the lineage. Many
aspects of pterosaur flight dynamics however, particularly those relating to the wing shape, joint mechanics and
a warping of the wing membrane during flapping flight, are still insufficiently understood.

Presented here are the latest experimental findings of the Karlsruhe based workgroup on pterosaur flight
dynamics. Using three-dimensionally preserved bones from a wide range of taxa the freedoms of movement
available to both the fore- and hind limb joints have been reconstructed. This allows for a more comprehensive
review of the wing/hind limb motion in both basal and derived pterosaurs that will be incorporated into future
experimental studies. In addition the initial experimental results of a wind tunnel, model-based approach to
understanding pterosaur flight dynamics is compared with those of a theoretical mathematical-based approach to
provide additional insights into how flight style evolved within the pterosaur lineage.

Key words: pterosaurs, flight dynamics, evolution, aerodynamics

36

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
DIET EVIDENCES ON VISEAN CRUSHING-TOOTH (COCHLIODUS)
FROM SOIGNIES (BELGIUM)

Fabrice FACK 1, Claire DERYCKE 2

1 6 carrière Rasson 59150 Wattrelos (France), fabrice.fack@wanadoo.fr
2 Université Lille 1, Laboratoire Géosystèmes (UMR 8157 CNRS),UFR des Sciences de la Terre - bâtiment
N5, 59655 Villeneuve d'Ascq cedex (France)& National Museum of History, Paleobiodiversity, UMR 5143.
Paris, claire.derycke@univ-lille1.fr

Begun thirty years ago (Scott et al., 2008), studies of dental microwear traces were essentially made on
extinct mammals. Most of these analyses of fossil forms concern anthropoids, primates, herbivorous and
carnivorous mammals (Goswani et al., 2005). Other but fewer treated amniotes like cynodont or dinosaur teeth
(Fiorillo, 1998 in Goswani et al., 2005). These traces could be related with diet and are considered more useful
than molar morphology. All these studies concern terrestrial forms practising mastication. Comparisons are
available for aquatic mammals too like odontocetes (Caldwell & Brown, 1964) and sirenians (Domning &
Beatty, 2007) but for the last, mainly on tusks, incisor teeth used for gathering food. Wear traces were examined
on grasping teeth of extant fishes (Purnell et al., 2006). Before, attempts were made to find wear traces on
conodonts (Donoghue & Purnell, 1999) considered by some authors as vertebrates. Presence of striae was
signalled in the oldest holocephalan crushing-tooth (Darras et al., 2008) and exposed, for the first time, in a true
palaeozoic fish tooth (Derycke, 2008). Contrary to multicuspidate teeth of chondrichthyans with a continuous
replacement of shedding teeth, crushing-teeth are considered to growth continuously and are more subject to
abrasion due to a slow replacement rate. The treatment of the prey even in water is comparable with what happen
on mammal molars. From scanning electron microscopy photos, measurements of striae have been made with
the software developed by Ungar Microware 4.02 even if today some authors advise to come back to low
magnification (Teaford et al., 2008). This software takes three measures for each stria: length, width, and
orientation. Two essential conclusions can be extracted from the observation of striation about the diet of
Cochliodus : 1) analysis of the tooth surface reveals two distinct families of oblique striae generated on a distal
tooth by a vertical movement of mastication, 2) distribution of feature orientations is less homogenous than seen
on a typical pointed tooth of a carnivore and striation length is smaller than exposed on a primate tooth under an
optical microscope. These observations show mastication of hard objects and could be due to the sediment
influence (Zangerl, 1981; Purnell et al., 2006) and/or to a durophageous diet. Concerning biomechanical
considerations, the hardness of apatite (5) for the predator teeth is higher than the calcite one (3) for the prey test.
Wear traces also depend on tissue density, on hardness increasing with the mineralization degree and on
microstructural orientation of fibers. For example, erosion is more rapid when fibers are parallel with the surface
and less rapid when perpendicular. Even soft abrasive particles cause erosion (Boyde, 1984).

BOYDE A., 1984. Dependence of rate of physical erosion on orientation and density in mineralised tissues. Anatomy and Embryology, 170 :
57-62.
CALDWELL, D. K. & BROWN D. H., 1964. Tooth wear as a correlate of described feeding behaviour by the killer whale, with notes on a
captive specimen. Bulletin Southern California Academy of Sciences, 63(3):128-140.
DARRAS, L., DERYCKE, C., BLIECK, A. & VACHARD, D., 2008. The oldest holocephalan (Chondrichthyes) from the Middle Devonian
of the Boulonnais (Pas-de-Calais, France). C. R. Palevol, 7 : 297-304.
DERYCKE, C., 2008. Comparisons between oldest chondrichthyan crushing-tooth and tooth with cusps. Journal of Vertebrate Paleontology,
28(3): 71A.
DOMNING, D.P & BEATTY, B.L., 2007. Use of Tusks in Feeding by Dugongid Sirenians : Observations and Tests of Hypotheses. The
Anatomical Record, 290 : 523-538.
DONOGHUE, P. C. J. & PURNEL, L M. A., 1999. Mammal-like occlusion in conodonts. Paleobiology, 25(1), pp. 5874.
GOSWANI, A., FLYNN, J.J., Ranivoharimanana, L., & Wyss A.R. 2005. Dental microwear in Triassic amniotes : implications for
paleoecology and masticatory mechanics. Journal of Vertebrate Paleontology, 25(2) : 320-329.
PURNELL, M. A., HART, P. J. B., BAINES, D. C., & BELL, M. A., 2006. Quantitative analysis of dental microwear in threespine
stickleback: a new approach to analysis of trophic ecology in aquatic vertebrates. Journal of Animal Ecology, 75, 967977.
SCOTT, R., SCHUBERT, B., GRINE, F., & TEAFORD, M., 2008. Low magnification microwear: question of precision and repeatability.
Journal of Vertebrate Paleontology, 28(3): 139A.
TEAFORD, M., GRINE, F., KAY, R., SCHUBERT, B., & UNGAR, P., 2008. Low magnification dental microwear : the problem of
postmortem artefacts. Journal of Vertebrate Paleontology, 28(3): 151A.
ZANGERL, R., 1981. Paleozoic Elasmobranchii Chondrichthyes. In: Schultze, H.P. (ed.), Handbook of Paleoichthyology, 3A, Stuttgart, 115
p.

Key words: dental microwear, Chondrichthyes, Visean, Soignies

37

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
PERMOCARBONIFEROUS AMNIOTES FROM FRANCE: A STATE OF
THE ART AND PERSPECTIVE

Jocelyn FALCONNET

UMR 5143 CNRS, Département Histoire de la Terre,
Muséum national d'Histoire naturelle, CP 38, 57 rue Cuvier, 75005 Paris, France, falconnet@mnhn.fr

The Paleozoic continental basins of France have yielded a rather diverse tetrapod fauna. They include
an amount of temnospondyls, of which many archegosaurids and eryopoids, but also a few lepospondyls and
amniotes. These amniotes except Casea rutena are especially poorly known, as they are commonly
represented by a single scrappy specimen, hence a complicated taxonomic history. Moreover, the recent
statements about their validity and affinities are based on outdated descriptions and figures published more than
eighty years ago. Consequently, a synthesis of systematical, geographical, and stratigraphical information
published or not is given for each of these taxa, as a basis for a future revision.
The Blanzy-Montceau Basin yielded the type specimen of the ophiacodontid Stereorachis blanziacensis
Langiaux, Parriat & Sotty, 1974, the sole amniote occurring from Stephanian strata in France. A close relative,
Stereorachis dominans Gaudry, 1880, is known from the Autunian deposits of the Autun Basin, along the basal
sphenacodonts Haptodus baylei Gaudry, 1886, and Callibrachion gaudryi Boule & Glangeaud, 1893, and an
indeterminate bolosaurid parareptile. A sphenacodontid of the same age, Neosaurus cynodus (Gervais, 1869)
Nopsca, 1923, was found in the Permian beds of the La Serre Massif Basin. The Lodève Basin, to the contrary,
produced remains of various Autunian amniotes, including the diapsid Aphelosaurus lutevensis Gervais, 1859,
and several partial remains belonging to an indeterminate caseid, to ?Stereorachis, as well as two new specimens
identified as an indeterminate sphenacodont and a new amniote of uncertain affinities. Besides, another
indeterminate caseid was found in the upper Permian series (`Saxonian') of this basin. The same goes for the
Rodez Basin, where two caseids were collected, the first being now Casea rutena and the second still
indeterminate. Finally, two indeterminate reptiles were produced by `Saxonian' deposits of the Saint-Affrique
Basin.
Among these taxa, several are now considered as nomen dubium (Haptodus baylei, Callibrachion,
Neosaurus), and one have to be assigned to a new genus (Casea rutena) because the current one appears to be
paraphyletic. Actually, most of these taxa must be redescribed adequately to assess their validity and then, if
valid (and possible), should be included in a phylogenetic analysis to understand their relationships with the
better known forms found in North America, and Europe.

Key words: Paleozoic, France, Amniota, synthesis

38

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
NEW DATA ON THE PALAEOBIOGEOGRAPHY OF TOARCIAN
(LOWER JURASSIC) ICHTHYOSAURS

Valentin FISCHER 1, Myette GUIOMAR 2, Pascal GODEFROIT 3

1 Unite de Paleontologie animale et humaine, University of Liege, Boulevard du Rectorat, 4000 Liege,
Belgium, val.fischer@student.ulg.ac.be
2 Reserve naturelle geologique de Haute Provence, montée Bernard Dellacasagrande 10, 04005 Digne les
bains, France, m.guiomar@resgeol04.org
3 Royal Belgian Institute of Natural Sciences, rue Vautier 29, 1000 Brussels, Belgium,
Pascal.Godefroit@naturalsciences.be

The Vocontian Basin of SE France was formed along the northwestern border of Tethys during Mesozoic
times. Mainly known for its rich ammonite fauna, this basin has also yielded Jurassic and Cretaceous ichthyosaur
fossils. The specimens discussed here were discovered in lower Toarcian limestone and marl successions in the
vicinity of Digne-les-Bains, High-Provence Alps. One of best-preserved specimens is identified as
Suevoleviathan sp., a relatively rare genus previously reported only in the German basins. The specimen is fairly
complete but crushed and embedded in limestone from the Falciferum ammonite zone. Because the skull is too
damaged to see bone sutures, assignation to this genus is mostly based on fin and coracoid morphology. Along
with this specimen, a premaxilla and paddle elements of Eurhinosaurus longirostris and possible
Stenopterygiidae centra were found in nearby black marls of the Exaratum ammonite subzone (Falciferum zone,
Lower Toarcian; see Floquet et al., 2003 for a detailed stratigraphy). These soft marls were deposited in anoxic
waters and are coeval with the Posidonia shales of southwest Germany, the Jet rock formation of northeast
England and Grandcourt shales of Luxembourg, which allows faunal comparisons between these basins. The
localities from the Vocontian Basin are closer to the Tethys than other sites where Toarcian identifiable
ichthyosaurs have been found in Europe. Nevertheless all identifiable specimens from the Vocontian Basin are
also ientified in other basins across Europe. It suggests that Toarcian ichthyosaurs had wide palaeogeographical
distributions and were highly mobile swimmers, perfectly adapted to open marine environments.

Key words: Ichthyosauria, Suevoleviathan, Palaeobiogeography, Lias, SE France

39

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
A NEW IGUANODON SPECIES (DINOSAURIA, ORNITHISCHIA)
FROM THE LOWER CRETACEOUS OF SORIA (SPAIN)

Carolina FUENTES VIDARTE 1, Manuel MEIJIDE CALVO 1, Federico MEIJIDE
FUENTES 2, Manuel MEIJIDE FUENTES 3

1 c/ Almazán 17, 2º C, E-42004 Soria, Spain, carolfuentes@ono.com
2 c/ Juan Sala de Pablo 8, 1º C, E-42003 Soria, España, fmeijidef@jazzfree.com
3 c/ Condes de Gómara 8, 3º B, E-42002 Soria, España

The Zorralbo site near Golmayo (Soria, Spain) belongs to the upper Hauterivian lower Barremian
Golmayo Formation (Clemente & Alonzo, 1990; Martín-Closas & Alonso-Millán, 1998). This locality has
yielded a diversified vertebrate assemblage: dinosaurs, a turtle and a "mesosuchian" crocodilian.
Dinosaurs are represented, in order of relative abundance, by ornithopods (Iguanodon), ankylosaurs
(Polacanthus), sauropods (basal member of Titanosauriformes) and indeterminate theropods (Fuentes Vidarte et
al., 2005).
The Iguanodon remains consist of more than 400 cranial, axial, and appendicular elements. Most of the
bones, if not all, belong to a single medium- to large-sized adult individual.
A detailed osteological study of the Soria material reveals significant differences with currently
dscribed Iguanodon material, suggesting that this new specimen would belong to a new Iguanodon species.

CLEMENTE, P. & ALONSO, A., 1990. Estratigrafía y sedimentología de las facies continentales del Cretácico inferior en el borde
meridional de la cuenca de los Cameros. Estudios Geológicos, 46: 257-276
FUENTES VIDARTE, C., MEIJIDE CALVO, M., MEIJIDE FUENTES, F. ,& MEIJIDE FUENTES, M., 2005. Fauna de vertebrados del
cretácico inferior del yacimiento de "Zorralbo" en Golmayo (Soria, España). Revista Española de Paleontología N. E., 10: 83-92.
MARTÍN-CLOSAS, C. & ALONSO-MILLÁN, A., 1998. Estratigrafía y bioestratigrafía (Charophyta) del Cretácico inferior en el sector
occidental de la Cuenca de cameros (Cordillera Ibérica). Revista Sociedad Geológica España, 11: 253-269.

Keywords: Iguanodon, Lower Cretaceous, Hauterivien-Barremien, Ornithopoda, Soria, Spain

40

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
CHELONIANS FROM THE LATE MIOCENE OF CHAD
(THE HOMINID SITE TM 266)

Géraldine GARCIA 1, Likius ANDOSSA 2, Hassan-Taisso MACKAYE 2,
Patrick VIGNAUD 1, Michel BRUNET 1-3

1IPHEP, Université de Poitiers, 40 Avenue du Recteur Pineau, 86022 Poitiers cedex, France
2Département de Paléontologie, Université de N'Djamena, BP 1117, N'Djamena, Chad
3Collège de France, Chaire de Paléontologie Humaine, 3 rue d'Ulm, 75231 Paris cedex 05, France

Fossil turtles from the Toros-Menalla localities (Late Miocene, Chad) are abundant and well diversified
with three families (Pelomedusidae, Testudinidae and Trionychidae) and at least four different species. The
chelonian material collected at TM 266, the site that has yielded the cranium of the earliest known hominid,
Sahelanthropus tchadensis (Brunet et al. 2002, 2005), includes remains belonging to the modern genus of
Cyclanorbinae (Trionychidae) and also one land tortoise (Testudinidae). The significant occurrence of aquatic
soft-shelled forms in association with terrestrial specimens is consistent with paleoenvironment reconstructions
of mosaic landscapes (Vignaud et al. 2002; Brunet et al. 2005, Le Fur et al. in press).

BRUNET, M. et al., 2005. New material of the earliest hominid from the Upper Miocene of Chad. Nature, 434: 752-755.
BRUNET, M. et al., 2002. A new hominid from the Upper Miocene of Chad, Central Africa. Nature, 418: 145-151.
LE FUR, S. et al., in press. The mammal assemblage of the hominid site TM 266 (Late Miocene, Chad Basin): ecological structure and
paleoenvironmental implications. Naturwissenchaften.
VIGNAUD, P. et al., 2002. Geology and Palaeontology of the Upper Miocene Toros-Menalla fossiliferous area, Djurab Desert, Northern
Chad. Nature, 418: 152-155.

Key words: chelonians, Chad, Late Miocene, palaeoenvironments

41

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
AMNIOTIC EGGSHELLS FROM THE HAlEG BASIN
(UPPER CRETACEOUS, ROMANIA)

Géraldine GARCIA 1, Pascal GODEFROIT 2, Thierry SMITH 2,
Jimmy VAN ITTERBEECK 3, Xavier VALENTIN 1, Vlad CODREA 4

1IPHEP, Université de Poitiers, 40 Avenue du Recteur Pineau, 86022 Poitiers cedex, France,
geraldine.garcia@univ-poitiers.fr
2 Department of Palaeontology, Royal Belgium Institute of Natural Sciences, rue Vautier 29, 1000 Brussels,
Belgium, Pascal.Godefroit@naturalsciences.be, Thierry.Smith @naturalsciences.be
3 Afdeling Historische Geologie, Katholieke Universiteit Leuven, Redingenstraat 16, B-3000, Leuven,
Belgium
4 Catedra de Geologie-Paleontologie, Universitatea Babe -Bolyai, Str. Koglniceanu 1, 3400 Cluj-Napoca,
Romania, codrea_vlad@yahoo.fr, vcodrea@bioge.ubbcluj.ro

Abundant eggs and eggshells were collected in the Upper Cretaceous deposits from the Haeg Basin
(Romania) during these last decades (Grigorescu et al.1994, Garcia et al. 2002, Codrea et al. 2002). Their
examination has revealed an important taxonomic diversity the different amniotic clades being represented. Five
egg types (testudoid, crocodiloid, geckonoid, dinosaurid and ratite) have been identified from the Romanian
main localities: Pui, Tu tea, Tote ti-baraj and Nla-Vad. Their study provides new data on paleogeographical
and paleoenvironment and new insight on reptilian diversity during the Campano-Maastrichtian.

CODREA, V et al., 2002. Dinosaur egg nests, mammals and other vertebrates from a new Maastrichtian site of the Hateg Basin (Romania).
Comptes-Rendus Palevol, 1: 173-180.
GARCIA, G. et al., 2002. Parataxonomic classification of eggshells from Pui in the Hateg Basin (Romania). 7th European workshop of
vertebrate palaeontology, Sibiu (Romania).
GRIGORESCU, D. et al., 1994. Late Maastrichtian dinosaur eggs from the Hateg Basin (Romania). In Carpenter, K., Hirsch, K.F. and
Horner, J.R (eds.) Dinosaur Eggs and Babies, Cambridge University Press, pp. 75-87.

Key words: eggshells, Haeg Basin, amniotes, Romania

42

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE DINOSAUR ASSEMBLAGE OF THE EL CASTELLAR
FORMATION (UPPER HAUTERIVIAN LOWERMOST BARREMIAN,
TERUEL, SPAIN)

José Manuel GASCA, Miguel MORENO-AZANZA, José Ignacio CANUDO

Grupo Aragosaurus, Paleontología, Facultad de Ciencias. Universidad de Zaragoza. Pedro Cerbuna 12,
50009 Zaragoza, Spain, gascajm@unizar.es, mmazanza@unizar.es, jicanudo@unizar.es

The El Castellar Formation is an Early Cretaceous synrift unit within the Wealden facies. It presents the
richest Mesozoic terrestrial vertebrate remains in the Iberian Peninsula as well as major dinosaur diversity.
Nearly one hundred vertebrate fossil sites are known in this formation. Most of these sites are situated in the
Galve sub-basin (Teruel, Spain), a 40-km-long, 20-km-wide, NNW-SSE-elongated basin. This is a small rift
basin, which formed part of the larger Cretacic Maestrazgo Basin. It is located in the eastern part of the Iberian
Chain (NE Spain).
The sedimentary record of the El Castellar Formation, which is Upper Hauterivian Lowermost
Barremian in age and around 50 to 150 m in thickness, is divided into two stages by a marlstone interval with
gypsum. Stage 1 (clays intercalated with sandstones and limestones) shows a great variety of subenvironments
and facies (alluvial, palustrine and lacustrine), whereas in stage 2 (marls and limestones) an extensive, shallow
carbonate lake developed. The greatest accumulation of vertebrate remains is located in the palustrine-lacustrine
deposits at the base of stage 2.
The dinosaur remains are frequently associated with abundant vertebrate, invertebrate and plant
microfossils such as charophyte utriculi and ostracods. In our present study we have integrated the research done
in the last 20 years and the exhaustive prospection of the formation that is now being carried out. The following
vertebrate groups are among those recognized: osteichthyes, chondrichthyes (hybodontid sharks), amphibians,
squamates, chelonians, neosuchian crocodiles, pterosaurs and mammals. They are represented by isolated teeth,
eggshell fragments, coprolites, scales, plates and fragmentary bones. The dinosaur remains from the El Castellar
Formation usually comprise disarticulated elements, but there are some exceptional cases, such as the
remarkable presence of the sauropod Aragosaurus ischiaticus Sanz, Buscalioni, Casanovas & Santafé, 1987.
Isolated teeth are the most common type of dinosaur remains, along with fragmentary vertebral centra and bone
splinters.
The main groups of Early Cretaceous dinosaurs are present in the El Castellar Formation. Sauropods are
represented by a high diversity of macronarians (Aragosaurus ischiaticus, Oplosaurus armatus, "Pleurocoelus"
valdensis, Euhelopodidae, Camarasauridae?). Theropods are represented by basal tetanurans (two species of
Baryonychinae, Allosauroidea?) and Maniraptoriformes (indet. A and B, "Paronychodontids", Maniraptora
indet. A and B, "Prodeinodon", Velociraptorinae). Ornithopods (Heterodontosauridae?, "Hypsilophodontidae",
at least two species of Iguanodontoidea) and thyreophorans (Ankylosauria, Stegosauria) make up the faunal list
derived from dinosaur skeletal remains.
The fragmentary nature of the fossil material from the dinosaurs of the El Castellar Formation makes it
difficult to make specific assignations at present. However, the association does bear witness to the unusually
high palaeobiodiversity (21 taxa) characteristic of this part of the Iberian Peninsula during the Hauterivian
Barremian transition. The studies of the vertical and lateral distribution of the taxa currently being undertaken
will permit us to establish whether there is a palaeoenvironmental or palaeoecological distribution of the
dinosaurs in the El Castellar Formation.

Key words: Lower Cretaceous, Iberian Basin, Teruel, dinosaur, faunal list

43

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
IGUANODON BERNISSARTENSIS FROM THE EARLY APTIAN OF
MORELLA (CASTELLÓN, SPAIN)

José Miguel GASULLA 1, José Luís SANZ 1, Francisco ORTEGA 2,
Fernando ESCASO 1,3

1 Unidad de Paleontología. Departamento de Biología. Facultad de Ciencias. Universidad Autónoma de
Madrid. Cantoblanco, 28049 Madrid. gasuviol@terra.es
2 Facultad de Ciencias. UNED. c/ Senda del Rey, 9. 28040 Madrid. fortega@ccia.uned.es
3 Museo de las Ciencias de Castilla-La Mancha. Plaza de la Merced 1, 16001, Cuenca.

Iguanodon is the most abundant dinosaur represented in the Lower Cretaceous outcrops of Morella
(Castellon, Spain). The first discoveries in the area are closely linked to the history of Spanish dinosaurology. In
fact, the presence of Iguanodon in Morellan deposits is part of the first written reference about dinosaurs in
Spain in the late nineteenth century (Vilanova, 1872, 1873). The available material is generally assigned to I.
bernissartensis from since the first monograph on Spanish dinosaurs (Sanz et al, 1982). At present, the
Iguanodon remains has increased significantly due to material from the "Cantera del Mas de la Parreta", a quarry
for the exploitation of clay in which they have identified several fossil-sites (Gasulla, 2005).

One of these fossil sites, the denominated CMP-5, had yielded a partial and disarticulated Iguanodon
skeleton composed by more than 130 cranial and postcranial elements and fragments. The CMP-5 place is
located in the lower part of the deep red clay package belonging to the "Arcillas de Morella" Formation. This
Formation is mainly constituted by terrigenous sediments that it is located at the beginning of an Early Aptian
depositional sequence, being part of the large mesozoic sedimentary basin of "El Maestrazgo".

The locality corresponds to a small channel included in a muddy deltaic plain. The Iguanodon remains
were found in a concentration deposit of hardly 15 m2. The fossils are placed without a preferential orientation in
a horizontal plane and with its long axis parallel to the plane of sedimentation. The elements are not in
anatomical connection and show a low dispersion. The most complete skeletal parts correspond to the cervical,
dorsal, sacral and antero-caudal regions of the axial skeleton, as well as pectoral and pelvic girdles. Other
remains are some mandibular elements and several anterior autopodial remains, being the most significants the
dental pieces and the ungual phalanx of the digit I (pollex).

It has been analyzed the phylogenetic position of the individual using the data matrix proposed by
Norman (2002) The three maximum parsimony trees obtained show that the Iguanodon of CMP-5 is the sister
taxon of I. bernissartensis. In the revised version of the taxonomic analysis of Iguanodon proposed by Paul
(2008), the checked characters in the CMP-5 specimen also coincide with the condition attributed to I.
bernissartensis. We therefore propose to assign the CMP-5 individual to I. bernissartensis, confirming the
distribution of this taxon in the Lower Cretaceous of the Iberian Peninsula.

GASULLA, J. M. 2005. Los dinosaurios de Morella (Castellón, España): historia de su investigación. Revista Española de Paleontología
n.e. X: 29-38.
NORMAN, D.B. 2002. On Asian ornithopods (Dinosauria: Ornithischia). 4. Probactrosaurus Rozhdestvensky, 1966. Zoological Journal of
the Linnean Society. 136: 113-144.
PAUL, G.S. 2008. A revised taxonomy of the iguanodont dinosaur genera and species. Cretaceous Research, 29: 192-216.
SANZ, J.L.; CASANOVAS, Mª. L., Y SANTAFE, J. Vte. 1982. Paleontología. In: Santafé, J.V., Casanovas, M.L., Sanz, J.L. y Calzada, S.
(1982): Geología y Paleontología (Dinosaurios) de las Capas rojas de Morella (Castellón, España). Diputación Provincial de
Castellón y Diputación de Barcelona: 69-169.
VILANOVA, J. 1872. Compendio de Geología. Madrid. 588 pp.
VILANOVA, J. 1873. Act. Soc. Esp. Hist. Nat., 2: 8.

Key words: Iguanodon, early Aptian, Morella (Spain)

44

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE DOMESTICATION OF THE DOG DURING THE UPPER
PALAEOLITHIC

Mietje GERMONPRÉ

Department of Palaeontology, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, 1000
Brussels, Belgium, Mietje.Germonpré@naturalsciences.be

The evolutionary origin of the dog from wolves is well established via morphological and genetic data.
Between 14,000 and 10,000 years ago dogs are known from Western Europe, Southern Europe, the Near East,
the Russian Plain and Kamchatka. Genetic results suggest a much older origin of dogs than indicated by
prehistoric finds (Vila et al., 1997). According to Lindblad-Toh et al. (2005), an ancient genetic bottleneck
accompanying the domestication of dogs occurred around 27,000 years ago. With this in mind, the low
frequency of recognised dog skulls in Upper Palaeolithic sites is somewhat surprising. In our opinion, it is likely
that a number of Palaeolithic dog remains have so far not been recognized. We conducted an osteometric
analysis on a number of fossil large canids with the aims of identifying and distinguishing Palaeolithic dogs from
fossil wolves. Based on multivariate techniques, we could show that the skull morphology of a number of
specimens is very similar and distinct from the morphology of wolves. These fossil skulls are interpreted as
originating from Palaeolithic dogs. The oldest dog in our data set, with an AMS age of c. 31,700 BP, is from the
Goyet cave (Belgium). The age of the skull implies that it could be derived from an Aurignacian occupation. We
therefore tentatively propose, pending further evidence that the domestication of the dog had already begun in
the Aurignacian. The Goyet skull is very similar to Epigravettian Eliseevich (Sablin and Khlopachev, 2002),
Mezin 5490 and Mezhirich dog skulls, which are about 18,000 years younger. Compared to fossil and recent
wolves, Palaeolithic dogs exhibit a shorter and broader snout and a relatively wide brain case, but a reduction in
carnassial tooth length can not be observed. Their skull size suggests that these first dogs were large animals.
They may have been used for helping with the tracking, hunting or transport of large, "ice-age" game
(Germonpré et al., in press).

GERMONPRÉ, M., SABLIN, M.V., STEVENS, R.E., HEDGES, R.E.M, HOFREITER, M., STILLER, M., DESPRÉS, V.R., in press.
Fossil dogs and wolves from Palaeolithic sites in Belgium, the Ukraine and Russia: osteometry, ancient DNA and stable isotopes.
Journal of Archaeological Science, DOI 10.1016/j.jas.2008.09.033.
LINDBLAD-TOH, K., et al., 2005. Genome sequence, comparative analysis and haplotype structure of the domestic dog. Nature,438: 803-
819.
SABLIN, M.V. & KHLOPACHEV, G.A., 2002. The earliest Ice Age dogs: evidence from Eliseevichi I. Current Anthropology: 43, 795-799.
VILÀ, C., SAVOLAINEN, P., MALDONADO, J.E., AMORIM, I.R., RICE, J.E., HONEYCUTT, R.L., CRANDALL, K.A., LUNDEBERG,
J., WAYNE, R.K., 1997. Multiple and ancient origins of the domestic dog. Science, 279: 1687-1689.

Key words: dogs, domestication, Aurignacian, Goyet

45

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
EARLY LAND PLANT EVOLUTION: A TREMENDOUS SUCCESS
STORY

Philippe GERRIENNE

University of Liège, Palaeobotany, Palynology and Micropalaeontology, Department of Geology,
B18, Sart Tilman, 4000 Liége, Belgium, p.gerrienne@ulg.ac.be

Plants most probably started to invade the land at the beginning of the Palaeozoic Era. The first fossil
evidence for land plants (Embryophytes) comes from dispersed spores: the earliest generally accepted reports are
from rocks of mid-Ordovician age (Llanvirn, 475 million years ago). The earliest body fragments of these plants
are parts of sporangia of late Ordovician age (Caradoc, 450 million years ago), and the oldest fertile axial land
plant fossil is reported from the Middle Silurian (Wenlock, 425 million years ago). From the Late Silurian
(Pridoli, 418 million years ago) onwards, land plant populations evolved from patchy stands of Cooksonia-type
plants (a few centimetres high, isotomously branched, with terminal sporangia) to world-wide distributed forests
dominated by Archaeopteris majestic trees at the end of the Devonian (Famennian, 360 million years ago) or
gigantic Lycophytes and Sphenophytes during the Carboniferous (359-299 million years ago). On their way to
the greening of the continents, plants evolved anisotomous branching, indeterminate growth, secondary tissues
including wood and bark, long-lived roots, leaves, heterospory and seeds. All those innovations allowed them to
increase size, to enhance propagule dispersion and efficiency, and to colonize almost all terrestrial habitats. This
had a global scale impact on geophysical and geochemical cycles that peaked during the late Devonian, when
Earth experienced one of the 5 major past extinction events. This talk focuses on the evolutionary history of key
characters such as secondary xylem and seed habit, and on their implication on land plant communities. It also
summarizes the phylogenetic relationships of all major Devonian plant lineages.

Key words: Embryophyte evolution, Silurian, Devonian, Carboniferous

46

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
EARLY MAMMALIAN BRAIN DIVERSITY: INSIGHTS FROM
MULTITUBERCULATES

Emmanuel GILISSEN 1, Thierry SMITH 2

1 Royal Museum for Central Africa, Department of African Zoology, Leuvensesteenweg 13, B-3080 Tervuren,
and Université Libre de Bruxelles, Laboratory of Histology and Neuropathology CP 620, B-1070 Brussels,
Emmanuel.Gilissen@africamuseum.be
2 Royal Belgian Institute of Natural Sciences, Department of Paleontology, 29 rue Vautier, B-1000 Brussels,
Thierry.Smith@naturalsciences.be

According to Kielan-Jaworowska (1986, 1997), two distinct types of morphology characterize the brains
of Mesozoic mammals. In some primitive mammals such as Triconodon and multituberculates, the vermis and
the paraflocculi are very large with no apparent cerebellar hemispheres and no dorsal midbrain exposure
(cryptomesencephalic type). In the other type of brain morphology (eumesencephalic type), the cerebral
hemispheres are actually well developed, the presence of cerebellar hemispheres is apparent, and there is a large
dorsal midbrain exposure. Kielan-Jaworowska & Lancaster (2004) subsequently suggested to abandon the terms
cryptomesencephalic and eumesencephalic brains in the light of a new interpretation of multituberculate and
eutriconodontan endocasts. In this new interpretation, the region described as the "vermis" is rather an
impression of the superior cistern covering both the midbrain and the cerebellar vermis itself. A cistern is an
expansion of the subarachnoid space containing cerebrospinal fluid. This interpretation suggests that the superior
cistern must have been large enough in eutriconodonts and multituberculates to press on the internal table of the
cranial cavity during development in order to create the triangular bulge evident on the endocasts. Kielan-
Jaworowska & Lancaster (2004) and Kielan-Jaworowska, Cifelli & Luo (2004) do not use the term vermis
anymore for the triangular bulge visible on multituberculate and eutriconodontan endocasts but replace it by
"superior cistern". In support of this new interpretation, previous observations suggest that a similar "enlarged
vermis" exists in extant species such as the marsupial koala, in which this enlargement is due to the expansion of
the superior cistern, which makes a bulging impression on the skull. Kielan-Jaworowska & Lancaster (2004)
further precise that the midbrain (tectum mesencephali) is dramatically exposed in the koala brain but is covered
on the endocast. In the current study, we assess this new interpretation of the endocranial cast of Mesozoic
mammals with new observations of the endocranial cast morphology in extant monotremes (Zaglossus,
Tachyglossus, Ornithorhynchus), marsupials (Macropus, Thylacinus, Protemnodon, Sarcophilus, Perameles,
Phascolomys, Dasyurus, Phascolarctos, Didelphis, Lutreolina, Caluromys, Chironectes, Marmosa, Metachirus),
Tenrecidae and Dobsonia praedatrix. In all genera, the cerebellar vermis is clearly visible on endocranial casts.
In koalas (genus Phascolarctos) the triangular space covering the midbrain between the posterior portions of the
cerebral hemispheres is evident but cannot be confused with the cerebellar vermis, clearly apparent between the
cerebellar hemispheres. The bulging aspect of the triangular space between the cerebral hemispheres is mainly
due to the presence of the venous transverse sinuses, located anterior to the vermis. An interpretation alternative
to vermis or superior cistern could also be given. In the megachiropteran Dobsonia praedatrix for instance, this
bulging structure is the pineal organ (Bhatnagar et al., 1990). It is very proeminent on D. praedatrix endocranial
cast and covers the anterior part of the cerebellar vermis. On the basis of our current observations, we suggest
returning to the previous terminology used for describing the endocasts of Mesozoic mammals. It clearly appears
that the cerebellar vermis always makes a clearly visible impression on the posterior part of the endocranial
casts.

BHATNAGAR, K.P., FRAHM, H.D., & STEPHAN, H., 1990. The megachiropteran pineal organ: a comparative morphological and
volumetric investigation with special emphasis on the remarkably large pineal of Dobsonia praedatrix. J. Anat., 168: 143-166.
KIELAN-JAWOROWSKA, Z., 1986. Brain evolution in Mesozoic mammals. Contributions to Geology, University of Wyoming, Special
Paper, 3: 21-34.
KIELAN-JAWOROWSKA, Z., 1997. Characters of multituberculates neglected in phylogenetic analyses of early mammals. Lethaia, 29:
249-266.
KIELAN-JAWOROWSKA, Z. & LANCASTER, T.E., 2004. A new reconstruction of multituberculate endocranial casts and
encephalization quotient of Kryptobaatar. Acta Palaeontol. Pol., 49 (2): 177188.
KIELAN-JAWOROWSKA, Z., CIFELLI, R.L., & LUO, Z.-X., 2004. Mammals from the Age of Dinosaurs. Origins, Evolution, and
Structure. Columbia University Press, New York, 648 p.

Key words: Mesozoic mammals, multituberculates, brain evolution, cerebellum, cistern

47

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE LAST POLAR DINOSAURS: HIGH DIVERSITY OF LATEST
CRETACEOUS ARCTIC DINOSAURS IN RUSSIA

Pascal GODEFROIT 1, Lina GOLOVNEVA 2, Sergei SCHEPETOV 2,
Géraldine GARCIA 3, Pavel ALEKSEEV 2

1 Royal Belgian Institute of Natural Sciences, Department of Palaeontology, rue Vautier 29, B 1 000
Bruxelles, Belgium, Pascal.Godefroit@naturalsciences.be
2 Komarov Botanical Institute, Russian Academy of Sciences, Prof. Popov street 2,
St. Petersburg 197 376, Russia, LinaGolovneva@yandex.ru
3 Université de Poitiers Faculté des Sciences, CNRS UMR 6046 IPHEP, Institut International de
Paléoprimatologie et Paléontologie Humaine, Evolution et Paléoenvironnements,
40 avenue du Recteur Pineau, 86 022 Poitiers Cedex, France, geraldine.garcia@univ-poitiers.fr

A Late Maastrichtian vertebrate microfossil assemblage discovered at Kakanaut in north-eastern Russia
reveals that dinosaurs were still highly diversified in Arctic regions just before the Cretaceous-Tertiary mass
extinction Event: small ornithopods, hadrosaurids, ankylosaurians, neoceratopsians, troodontids, dromaeosaurids
and tyrannosaurids. Dinosaur eggshell fragments, belonging to hadrosaurids and non-avian theropods, indicate
that at least several latest Cretaceous taxa could reproduce in polar regions and were probably year-round
residents of high latitudes. Palaeobotanical data suggest that these polar dinosaurs lived in a temperate climate
(mean annual temperature about 10°), but the climate was apparently too cold for amphibians and ectothermic
reptiles. The high diversity of Late Maastrichtian dinosaurs in high latitudes, where ectotherms are absent,
invalidates hypotheses according to which dinosaur extinction was a result of temperature decline, caused or not
by the Chicxulub impact.

Key words: dinosaurs, Late Maastrichtian, polar regions, Russia

48

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
A NEW `MIDDLE' CRETACEOUS VERTEBRATE LOCALITY FROM
JILIN PROVINCE, NORTH-EASTERN CHINA

Pascal GODEFROIT 1, JIN Liyong 2, CHEN Jun 2, Richard BUTLER 3

1 Royal Belgian Institute of Natural Sciences, Department of Palaeontology, rue Vautier 29,
B 1 000 Bruxelles, Belgium, Pascal.Godefroit@naturalsciences.be
2 Jilin University Geological Museum, Chaoyang Campus, 6 Ximinzhu Street, Changchun, Jilin Province
130062, People's Republic of China, cj@jlu.edu.cn
3 Department of Palaeontology, The Natural History Museum, Cromwell Road,
London, SW7 5BD, United Kingdom, R.Butler@nhm.ac.uk

The Quantou Formation (late Early Cretaceous or early Late Cretaceous) is located along the eastern
margin of the Songliao Basin in Jilin Province, P.R. China. In 2000 and 2002, field parties were organized in a
manmade quarry, close to Gongzhuling City. Many vertebrate fossils were collected from a single horizon.

The most complete dinosaur material collected from the Quantou Formation is of the small
ornithischian Changchunsaurus parvus, which is represented by a well-preserved complete skull and partial
postcranial skeleton, as well as additional referred cranial and postcranial material. Changchunsaurus parvus
was originally briefly described by Zan et al. (2005), who recognised that this taxon possesses an unusual
combination of plesiomorphic and derived character states. New study of the type material reveals that
Changchunsaurus shares synapomorphies with the Chinese Early Cretaceous cerapodan Jeholosaurus, and it is
likely that they are sister taxa; moreover, Changchunsaurus possesses characters that suggest possible affinities
with marginocephalians (Ceratopsia + Pachycephalosauria).

A left maxilla and a right dentary can be attributed to a new basal neoceratopsian dinosaur (Jin et al., in
press). This new taxon differs from other basal neoceratopsians by its deep dentary ramus, the steeply-inclined
ventral predentary facet of its dentary, its heterogeneous dentary crowns, and by the denticles and secondary
ridges asymmetrically distributed on either side of the primary ridge on its dentary teeth. With Auroraceratops,
the new taxon from Gongzhuling represents one of the most derived non-coronosaurian neoceratopsians.

Isolated and fragmentary material of iguanodontian dinosaurs have also been discovered in this locality
(Chen et al., 2008). Dromaosaurids are represented by a huge raptorial sickle claw, still in articulation with a
small pedal giglymoid second phalanx, and by isolated teeth. Other theropods and crocodiles are also represented
by isolated teeth. Dinosaur eggs are abundant and diversified, but have not been studied yet.

Two incomplete mammalian dentaries represent a new taxon, named Zhangolestes jilinensis, referable
to the eutherian family Zalambdalestidae (Zan et al. 2006). A complete skull probably represents another new
genus of eutherian mammal.

The Quantou Formation fauna therefore has great potential for improving our understanding of the
fundamental reorganisation of Early - `middle' Cretaceous terrestrial ecosystems, which has been termed the
Cretaceous Terrestrial Revolution (Lloyd et al. 2008).

CHEN, J., BUTLER, R. J., & JIN, L.-Y., 2008. New material of large-bodied ornithischian dinosaurs, including an iguanodontian
ornithopod, from the Quantou Formation (middle Cretaceous: AptianCenomanian) of Jilin Province, northeastern China. Neues
Jahrbuch für Geologie und Paläontologie, Abhandlungen, 248: 309314.
JIN, L.-Y., CHEN, J., ZAN, S.-Q., & GODEFROIT, P., in press. A new basal neoceratopsian dinosaur from the `middle' Cretaceous of Jilin
Province, China. Acta Geologica Sinica.
LLOYD, G. T., DAVIS, K. E., PISANI, D., TARVER, J. E., RUTA, M., SAKAMOTO, M., HONE, D. E., JENNINGS, R., & BENTON,
M.J., 2008. Dinosaurs and the Cretaceous Terrestrial Revolution. Proceedings of the Royal Society B, Biological Sciences, 275:
24832490.
ZAN, S.-Q., CHEN, J., JIN, L.-Y, & LI., T., 2005. A primitive ornithopod from the Early Cretaceous Quantou Formation of central Jilin,
China. Vertebrata PalAsiatica, 43: 182193.
ZAN, S., WOOD, C. B., ROUGIER, G. W., JIN, L.-Y., CHEN, J., & SCHAFF, C. R., 2006. A new "middle" Cretaceous zalambdalestid
mammal, from a new locality in Jilin Province, northeastern China. Journal of the Paleontological Society of Korea, 22: 153172.

Key words: `middle' Cretaceous, north-eastern China, terrestrial ecosystem, dinosaurs, zalambdalestid eutherian
mammals

49

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
FROM PAST COLLECTIONS OF BERNISSART TO RECENT
COLLECTIONS OF HAUTRAGE: NEW TAXONOMICAL AND
ENVIRONMENTAL INSIGHTS

Bernard GOMEZ 1, Johan YANS 2, Thomas GILLOT 1, Paul SPAGNA 3, Clément
COIFFARD 4, Véronique DAVIERO-GOMEZ 1

1 CNRS-UMR 5125 Paléoenvironnements et Paléosphère, Université Lyon 1 (Claude Bernard), Campus La
Doua, Paléobotanique, Bât. Charles Darwin A, 7, rue Dubois, F-69622 VILLEURBANNE CEDEX,
FRANCE, bernard.gomez@univ-lyon1.fr, tom6446@hotmail.fr, daviero@univ-lyon1.fr
2 Facultés Universitaires Notre-Dame de la Paix, Département de Géologie, 61 rue de Bruxelles, 5000 Namur,
Belgium, johan.yans@fundp.ac.be
3 Faculté Polytechnique de Mons, Service de Géologie Fondamentale et appliqué, 9 rue de Houdain, 7000
Mons, Belgium; Paul.Spagna@fpms.ac.be
4 UMR 5143 du CNRS, Paléobiodiversité, systématique, évolution des embryophytes et Laboratoire de
paléobotanique et paléoécologie, université Pierre-et-Marie-Curie, département « Histoire de la Terre »,
MNHN, bâtiment de géologie, CP48, 57, rue Cuvier, 75231 Paris cedex 05, France, clement.coiffard@ens-
lyon.org

Previous studies from the Wealden facies of Belgium established the high diversity of fossil plants
(Bommer, 1892; Seward, 1900). However, except for the fern Weichselia reticulata (Alvin, 1968, 1971; Hill,
1990) and conifers and their allies (Harris, 1953; Alvin, 1953, 1957, 1960), most are misidentified, invalid or
poorly studied genera or species. The collections of the Royal Belgian Institute for Natural Sciences store figured
and type specimens, as well as hundreds of others in so numerous drawers. The taxonomy of specimens in
collections has been re-evaluated during the tenure of SYNTHESIS Project fellowship in January 2008. We have
particularly questioned whether the meso- and megafossil plants are actually exhaustively listed, and especially
for each locality. In parallel, we have sampled and are processing specimens from the Barremian-Aptian clay pit
of Danube-Bouchon's Hautrage quarry (Mons Basin, Belgium), which is coeval from the coal mine Sainte-Barbe
of Bernissart. The present paper compares both plant assemblages, and also gives new perspectives on taxonomy
(angiosperm leaves?), palaeoecology and palaeoenvironment (wildfires?).

ALVIN, K.L., 1953. Three Abietaceous cones from the Wealden of Belgium. Mémoires de l'Institut royal des Sciences naturelles de
Belgique, 125: 3-42.
ALVIN, K.L., 1957. On the two cones Pseudoaraucaria heeri (Coemans) n. comb. and Pityostrobus villerotensis n.sp. from the Wealden of
Belgium. Mémoires de l'Institut royal des Sciences naturelles de Belgique, 135: 3-27.
ALVIN, K.L., 1960. Further conifers of the Pinaceae from the Wealden Formation of Belgium. Mémoires de l'Institut royal des Sciences
naturelles de Belgique, 146: 3-39 (10 pls).
ALVIN, K.L., 1968. The spore-bearing organs of the Cretaceous fern Weichselia Stiehler. Journal of the Linnean Society (Botany), 61 (384):
87-92.
ALVIN, K.L., 1971. Weichselia reticulata (Stokes et Webb) Fontaine from the Wealden of Belgium. Mémoires de l'Institut royal des
Sciences naturelles de Belgique, 166: 1-33.
BOMMER, Ch., 1892. Nouveau gite de végétaux découvert dans l'argile wealdienne de Bracquegnies (Hainaut). Bulletin de la Société belge
de Géologie, de Paléontologie et d'Hydrologie, 6: 160-161.
HARRIS, T.M., 1953. Conifers of the Taxodiaceae from the Wealden Formation of Belgium. Mémoires de l'Institut royal des Sciences
naturelles de Belgique, 126: 3-43.
HILL, C.R., 1990. Scanning electron microscopy in palaeobotany. In: Claughter, D. (ed.), Scanning Electron Microscopy in Taxonomy and
Functional Morphology, Systematics Association Special Volume, 41, Clarendon Press, Oxford, pp. 193-234.
SEWARD, A.C., 1900. La flore wealdienne de Bernissart. Mémoires du Musée royal d'Histoire naturelle de Belgique, 1: 1-39.

Key words: Plant mesofossils, Bernissart, Hautrage, palaeoecology, palaeoenvironments

50

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
ZALMOXES, RHABDODON AND RHABDODONTIDAE:
HOW MANY GENERA AND HOW MANY SPECIES?

Florent GOUSSARD

Muséum National d'Histoire Naturelle, Département Histoire de la Terre, UMR 5143 CR2P CNRS,
8 Rue Buffon 75005 Paris, goussard@mnhn.fr

A new phylogenetic analysis of basal Euornithopoda calls into question the validity of the genus Zalmoxes
Weishampel et al., 2003 based on Transylvanian remains precedently referred to the genus Rhabdodon. The
revision of the available material permits to discuss the distribution of some characters ambiguous to date.
Consequently, the genus Zalmoxes Weishampel et al., 2003 is considered as junior synonym of the genus
Rhabdodon Matheron, 1869. This new study also confirms a basal position of Rhabdodon as sister-group of
Iguanodontia. Assumptions of sexual dimorphism and insular dwarfism are reconsidered in the issue of the
morphological variability of the genus Rhabdodon, leading to consider the two Romanian species (Zalmoxes
robustus and Zalmoxes shqiperorum, after Weishampel et al. 2003) like two potential sexual morphs of
Rhabdodon robustus.

Key words: Rhabdodon, Zalmoxes, Rhabdodontidae, systematics

51

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
REMORAS (TELEOSTEI, ECHENEIDAE) FROM THE OLIGOCENE
OF THE WEST CARPATHIANS (CZECH REPUBLIC)

Ruzena GREGOROVA

Department of Geology and Paleontology, Moravian Museum, Zelny trh 6, 659 37 Brno, Czech Republic,
rgregorova@mzm.cz

Remoras, or suckerfishes, are elongate brown fish; they grow up to 3090 centimeters long and their
distinctive first dorsal fin takes the form of a modified oval sucker-like organ with slat-like structures that open
and close to create suction and take a firm hold against the skin of larger marine animals.
There are 4 extant genera Echeneis, Phtheirichthys, Remora, Remorina, and 8 species. Remoras are
primarily tropical open-ocean dwellers, occasionally found in temperate or coastal waters if they have attached
to large fish that have wandered into these areas.
The fossil records of remoras are spare and up to now they are known only from the Tertiary sediments of
Paratethys (Carpathians, Caucasus) and Tethys (Glarus, Switzerland). The oldest record comes from the famous
Glarus locality in Switzerland, where Wettstein (1886) described a new Oligocene species Echeneis glaronensis.
Later this fossil representative was assigned to the genus Opistohmyzon (Cope 1889) and Berg (1958) placed it
into its own family Opisthomyzonidae. From the Oligocene of the Menilitic Formation of the Poland part of the
Carpathians Szajnocha (1926) described Echeneis carpathica (Wadovice locality). Recently this species was
recorded from eight other localities from the Menilitic Formation in Poland (Jerzmanska & Swidnicka, 2003).
Daniltshenko (1958) recorded Echeneis urupensis from the Miocene of North Caucasus. Micklich (1998)
described one specimen of Echeneidae as a probable representative of the genus Echeneis from the Frauenweiler
locality (Rupelian, Germany). Brzobohatý et al. (1974) mentioned Opisthomyzon from the Egerian locality of
Krumvi (Czech Republic).
The new fossil material of remoras from Moravia (Czech Republic) comes from the Sitboice beds
upper part of the Menilitic Formation, Nannoplankton Zone NP 23. A paper of O'Toole (2002) was used for the
osteological and comparative analyses of the fossil remnants that helped for the taxonomic determination.
The remoras from the two Moravian localities are preliminary attributed to the genera Remorina (2
specimens, Bystice/Olsí locality) and Remora (4 specimens, Litencice locality).
The following characters allow referring the fossil specimens to the genus Remorina: 25 26 vertebrae;
the neural spine on the preural centrum 3 is short, extending only to the anterior edge of the ural centrum;
number of the dorsal-fin rays = 23, number of anal fin-rays = 23. The following characters allow referring the
fossil specimens to the genus Remora: number of anal fin-rays =26; number of dorsal fin-rays > 26; strongly
curved teeth on dentary; the neural spine on the preural centrum 3 extends posteriorly behind the hypurapophysis
or extends between the anterior edge of the ural centrum and the hypurapophysis.
The anatomical study and phylogenetic analysis will include characters observed in other Tertiary fossil
material in order to understand the evolution and paleogeographical distribution of the family Echeneidae in the
region of the Paratethys.

Key words: Teleostei, Echeneidae, West Carpathians, Tertiary

52

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
DESCRIPTION OF THE MASTICATORY APPARATUS OF
LAONASTES AENIGMAMUS (RODENTIA, DIATOMYIDAE), NEW
INSIGHTS INTO THE EVOLUTION OF HYSTRICOGNATHY

Lionel HAUTIER 1, Soonchan SAKSIRI 2

1 Laboratoire de Paléontologie, Institut des Sciences de l'Evolution, Case Courrier 64, Université Montpellier
2, Place Eugène Bataillon, F-34095, Montpellier cedex 05, France. Emai : hautier@isem.univ-montp2.fr;
2 Department of Biology, Faculty of Science, Mahasarakham University; Tambon Khamriang Kantarawichai
district, 44150 Mahasarakham, Thailand.

Rodents have undergone an astonishing adaptive radiation through the Cenozoic that led them to
represent the most speciose order of mammals. Considering the direction of the angular process of the mandible
relative to the plane of the incisors, Rodentia were commonly divided into two suborders: Sciurognathi and
Hystricognathi. The living Laotian rock rat Laonastes aenigmamus, recently discovered (Jenkins et al. 2005) in
the Lao People's Democratic Republic (Lao PDR, Thakhek district), has first been considered as the sole
member of a new hystricognathous family Laonastidae. However, a reexamination of the specimens (Dawson et
al. 2006) has shown that this species could represent a surviving member of the extinct family Diatomyidae
among the "ctenodactyloid rodents", i.e. a sciurognathous family. More recently, molecular analyses (Huchon et
al. 2007) unambiguously confirmed the paleontological view in demonstrating that L. aenigmamus is the sister
group of Ctenodactylidae (within the monophyletic Ctenohystrica). Here, we provide the first complete
description of the masticatory apparatus of Laonastes aenigmamus. It offers a rare opportunity to study an
original morphological combination among Ctenohystrica and to investigate evolution of the masticatory
apparatus of hystricognathous rodents as a result. The gross anatomy and internal architecture of the jaw
musculature has been studied using dissections. Striking convergences have occurred during the evolution of
Diatomyidae and L. aenigmamus presents a unique combination of myological and osteological features that
corresponds to a mixture of sciurognathous and hystricognathous characters. This investigation lead to discuss
on hystricognathy (Tullberg, 1899), which is sometimes confused with sciurognathy, and on its phylogenetical
impact. These results bring new insights into the evolution of hystricognathy and have profound implications for
the interpretation of the early hystricognath rodents fossil record.

Key words: masticatory muscles, myology, Diatomyidae, hystricognathy

53

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
DIVERSITY AND DISPARITY IN SAUROPOD DINOSAURS

Christophe HENDRICKX 1, Steve BRUSATTE 2, Mark YOUNG 1, Emily RAYFIELD 1,
Marcello RUTA 1, Paul BARRETT 3

1 Department of Earth Sciences, University of Bristol, Bristol BS8 1RJ, UK, corresponding authors
christophe.hendrickx@hotmail.com
2 Division of Paleontology, American Museum of Natural History, New York, NY 10024, USA,
brusatte@gmail.com
3 Department of Palaeontology, The Natural History Museum Cromwell Road, London, SW7 5BD, UK,
p.barrett@nhm.ac.uk

A new aspect of sauropod history has been studied by analyzing the morphological variation of this
successful clade through time. Morphological diversification, when compared to taxonomic diversity, can reveal
macroevolutionary processes that shaped the history of a clade. This allows investigation into patterns of
morphological evolution. Results indicate that sauropod taxonomic richness and morphological variety increased
together during the initial diversification of the clade. This pattern suggests continued diffusion through
morphospace. The sauropod clade shows maximal morphological and taxonomic diversity in the Late Jurassic,
suggesting some degree of common control on diversification patterns in this group. A decline in taxonomic
richness in sauropods after the Jurassic/Cretaceous boundary is not accompanied by a corresponding reduction in
morphological diversity, suggesting non-selective extinction. Finally, an increase in taxonomic and
morphological diversity through the end of the Cretaceous corresponds to the radiation of titanosaurs.

Key words: Dinosauria, Sauropoda, Diversity, Disparity

54

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
MORPHOFUNCTIONAL ANALYSIS OF SPINOSAURID QUADRATES

Christophe HENDRICKX 1, Eric BUFFETAUT 2

1Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road,
Bristol BS8 1RJ, UK, christophe.hendrickx@hotmail.com
2CNRS (UMR 8538 Laboratoire de géologie de l'École Normale Supérieure), 16 cour du Liégat,
75013 Paris, France, eric.buffetaut@wanadoo.fr

Five quadrates from the Early Cenomanian of the Kem Kem area (Morocco) are determined to be from
juvenile and adult spinosaurids. Their morphology indicates two morphotypes and reveals the presence of two
different taxa of spinosaurids in this site. Morphofunctional analysis of quadrate bones and their mandibular
condyles has showed that the mechanics of the lower jaw of spinosaurids was specialized. The posterior parts of
the mandible displaced laterally when the jaw moved downwards thanks to a helicoidal shape of the mandibular
articulation of the quadrate. Such a lateral movement of the rami was also possible thanks to a weak and short
mandibular symphysis and allowed the pharynx to be widened. This jaw mechanic is present in some
ornithocheiroid pterosaurs and living pelecanid birds, which are both adapted to piscivory and to swallowing
large fish. Spinosaurids which were engaged in at least a partially piscivorous lifestyle were able to consume
large fish and may have fed occasionally other prey such as pterosaurs and juvenile dinosaurs.

Key words: Dinosauria, Spinosauridae, Quadrate, Cenomanian, Morocco

55

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
CONTRIBUTION TO THE EVOLUTIONARY HISTORY OF THE
BEAKED WHALES

Ella HOCH

Gram Museum of Palaeontology, DK-6510 Gram, Denmark, ella.hoch@mail.dk

Whales, Cetacea, evolved from terrestrial raoellid artiodactyls in warm southern Asia in the Early
Eocene. The pakicetids, the oldest known whales, approximately 50 Ma, had the general form of large dogs with
long snouts and long tails and were waders in shallow rivers and lakes (Thewissen, 2008a,b). In the later part of
the Eocene the basilosaurid whales were tropical cetaceans of a fully aquatic grade (Gingerich, 2005).
Archaeoceti adapted to life in water during the Eocene seemingly without acquiring food-gathering or diving
capabilities much above crocodilian states. Neoceti, comprising Mysticeti, the baleen whales, and Odontoceti,
the toothed whales, are derived from the dorudontine basilosaurids. The credible times of divergence
of Mysticeti and Odontoceti are calculated at near the Eocene/Oligocene boundary (Gingerich, 2005). The
process of change from archaeocetism to odontocetism may well have been spurred by cranial telescoping
initiating the evolution of structures and capacities for sophisticated sonar (echolocation), a character of the
odontocetes. Encephalization belongs to this complex of specializations, with the acoustic sense dominant.

Advanced sonar and complex body adaptations gave access to food webs far below the photic zone in
the global waters. Beaked whales and sperm whales among the odontocetes include the deepest-diving
mammals, and possibly the deepest-diving tetrapods through all ages. Some species may habitually descend to
depths of 1-2 kilometres (Physeter catodon may go deeper), feeding under water pressures of 100-200
atmospheres, yet breathing at the water's surface. Similarities between beaked whales and sperm whales may
reflect evolutionary convergence rather than being indicators of close relationship, a view contested by some
researchers. As it is illustrated by Bianucci & Landini (2007) and supported by other authors, the sperm whale
lineage, comprising Physeteridae, Kogiidae and extinct physeteroids, form an early, isolated evolutionary
branch, and the beaked whale lineage, Ziphiidae, with a fossil record not older than Burdigalian or late Early
Miocene, groups with the delphinoids and the platanistoids, and may have evolved from Early Miocene
eurhinodelphinids (Lambert, 2005). Extant ziphiids constitute the second largest odontocete family, after the
delphinids, counted in number of species; and Berardius bairdii, the northern giant bottle-nosed whale, is the
second largest living odontocete. Six genera are recognized in Recent time, which is slightly below the assumed
generic diversity of the ziphiids in Tortonian or early Late Miocene time (e.g. Lindberg & Pyenson, 2006).
Miocene ziphiids became known from the excavations of the docks at Antwerp mentioned with appreciation
by G. Cuvier in 1812 for the collections of fossils made by enlightened engineers. These ziphiids had lived in the
North Sea when it was open to the oceans only in northern direction, between Scotland and Norway, and the
regional climate tended towards cool temperate. Other Miocene ziphiids have been described on fossils from the
North Atlantic, the South Atlantic and the Mediterranean realms. In all cases the fossils were parts of skulls or of
lower jaws. A new extinct ziphiid, represented by associated skull parts, lower jaw, teeth and parts of the anterior
body skeleton from Tortonian shallow water deposits of the eastern North Sea, the Gram Formation, now
exposed on land in south-western Denmark, will be presented here. Darwin's famous "descent with
modifications" enounced change (of organisms) through time. It indicated no direction of the changes.
Circumstances and surroundings, including other living beings, would influence the direction of evolution, and
the changes in complexity of the organisms would be summarized by analysts in grades of specialization. The
new, approximately 8 million years old, ziphiid is studied with an aim of evaluating its grade of specialization
relative to those of the extant ziphiids.

BIANUCCI, G. & LANDINI, W., 2007. Fossil history. Pp 35-93 in D.L. Miller (ed.), Reproductive Biology and Phylogeny of Cetacea:
Whales, Dolphins and Porpoises. Science Publishers, Enfield, Jersey, Plymouth.
CUVIER, G., 1812. Recherches sur les ossemens fossiles de quadrupèdes Vol. 1, Avertissement, i-vi., Paris.
GINGERICH, P.D., 2005. Cetacea. Pp 234-252 in K.D. Rose and J.D. Archibald (eds), The Rise of Placental Mammals. The Johns Hopkins
University Press.
LAMBERT, O., 2005. Systematics and phylogeny of the fossil beaked whales Ziphirostrum du Bus, 1868 and Choneziphius Duvernoy, 1851
(Mammalia, Cetacea, Odontoceti), from the Neogene of Antwerp (North of Belgium). Geodiversitas 27, 443-497.
LINDBERG, D.R. & PYENSON, N.D., 2006. Evolutionary patterns in Cetacea. Fishing up prey size through deep time. Pp 67-81 in J.A.
Estes et al. (eds), Whales, Whaling, and Ocean Ecosystems. University of California Press.
THEWISSEN, J.G.M., 2008a. The origin of whales: Collecting fossils in the Himalayas. Fifth Conference on Secondary Adaptation of
Tetrapods to Life in Water, Abstracts, 72-73. Tokyo. ISBN 978-4-87803-024-6.
THEWISSEN, J.G.M., 2008b. Raoellids and archaeocetes: The first ten million years of whale evolution. Fifth Conference on Secondary
Adaptation of Tetrapods to Life in Water, Abstracts, 74. Tokyo. ISBN 978-4-87803-024-6.

Key words: Cetacea, Ziphiidae, Denmark, Gram Formation, Tortonian

56

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
DRAGONS (URSUS SPELAEUS) BY JOHAN PATERSON HAIN (1615 -
1675) FROM PIENINY (SLOVAKIA)

Zuzana KREMPASKÁ

The Museum of Spis in Spisská Nová Ves, Letná 50, 052 01 Spisská Nová Ves, Slovakia,
z.krempaska@gmail.com

The archive records about "Dragon Caves" from Slovakia are dated back to 17th century. Johan Paterson
Hain (1615 1675), a physician from Presov was the first who attempted to study the dargon bones from the
Aksamitka cave near the village of Haligovce, Pieniny (Slovakia). In 1672 he published his findings in
Miscellanea Curiosa Medico Physica Academiae Naturae Curiosorum sive Ephemeridum medico
physicarum Germanicarum. In the article entitled De Draconum Carpaticorum Cavernis he described and drew
the outline of the bones. Here he also mentions the healing effect of the bones and the appearance of the moon
milk (Album Nihil). Daniel Fischer (1695 1746), a physician from Kezmarok who followed the footsteps of
J.P.Hain, had reconstructed the skeleton of the "dragon" out of the bones from Aksamitka. He donated his
"masterpiece" to the emperor Leopold I.

The 330 m long Aksamitka cave (750m above the sea level) is situated in Pieniny National park
(Slovakia). Unfortunately, it is not opened to public.

Key words: Johan Paterson Hain (1615 1675), the dragon caves, Aksamitka cave in Pieniny (Slovakia)

57

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
NEW STEM SPERM WHALES FROM THE MIOCENE OF PERU

Olivier LAMBERT 1, Giovanni BIANUCCI 2, Christian de MUIZON 3,
Alton C. DOOLEY, Jr 4

1 Institut royal des Sciences naturelles de Belgique, Département de Paléontologie, rue Vautier, 29, B-1000
Bruxelles, Belgium, Olivier.Lambert@naturalsciences.be
2 Dipartimento di Scienze della Terra, Università di Pisa, via S.Maria, 53, I-56126 Pisa, Italy,
bianucci@dst.unipi.it
3 Muséum National d'Histoire Naturelle, UMR 5143 (Paléobiodiversité et Paléoenvironnements), rue Cuvier,
57, CP 38, F-75005 Paris, France, muizon@mnhn.fr
4 Virginia Museum of Natural History, Starling Avenue, 21, VA-24112 Martinsville,Virginia, USA
alton.dooley@vmnh.virginia.gov

In both modern sperm whale genera (Cetacea, Odontoceti, Physeteroidea), the huge Physeter and the
small Kogia, the upper jaw is edentulous. The main food item is cephalopods, which are swallowed by suction.
Physeter, which retains elongated jaws, sucks the squid directly into the oropharyngeal opening, whereas the
extremely shortened snout of Kogia makes easier the process of oral suction. In these two taxa the cranium is
highly modified, excavated by the supracranial basin, a deep dorsal fossa containing the spermaceti organ and
other structures associated to the production and transmission of the sounds. In Physeter, this basin extends far
anteriorly on the rostrum, corresponding to a large spermaceti organ giving to the head a typical barrel shape.

The oldest physeteroids are fragmentary remains dating from the Late Oligocene. Poorly diagnostic
isolated physeteroid teeth are relatively common in Miocene sediments, but specimens including well-preserved
cranial elements are rare. We report here two nicely preserved new skulls from the Miocene part of the Pisco
Formation, south coast of Peru. Both skulls are associated with the mandible and teeth. The first specimen, with
part of the hyoid bones and some vertebrae preserved, comes from the locality of Cerro la Bruja (ClB) and is
dated from the late Middle to early Late Miocene (~11-12 Ma). The second specimen, attributed to the new
genus and species Acrophyseter deinodon, was found in a latest Miocene layer (~ 6 Ma) in the locality of Sud-
Sacaco.

These two taxa share a moderate skull size (cranium length = 460mm for the ClB specimen, lacking the
anterior portion of the rostrum; condylobasal length < 840mm for the almost complete skull of the holotype of A.
deinodon). In A. deinodon each upper tooth row bears 12 teeth, with 13 in each lower row. The ClB specimen
has both maxillary and mandibular teeth as well, but the total tooth count is unknown. Upper and lower teeth are
robust, with a swollen root, and separated from each other by short interalveolar spaces. In both taxa the rounded
temporal fossa is considerably proportionally larger than in modern sperm whales, suggesting a massive jaw
musculature and a powerful bite. On the cranium, the supracranial basin is deep, with a high posterior wall. The
basin is especially wide on the right side compared to most other physeteroids, overhanging the right orbit and
antorbital notch. However, the morphology of the maxillae and premaxillae at the rostrum base indicate that the
basin is limited to the cranium, meaning that the spermaceti organ did not invade the dorsal surface of the
rostrum.

A preliminary phylogenetic analysis, only including A. deinodon, identified the latter as a stem
physeteroid, more basal than the clade including Physeter (family Physeteridae), Kogia (family Kogiidae), their
last common ancestor, and all its descendants.

The slightly smaller and stratigraphically younger A. deinodon has a shorter, more pointed rostrum than
the ClB specimen. Furthermore its mandible is more upturned; the zygomatic process of the squamosal is more
pointed; the temporal fossa is more posterodorsally extended; and the coronoid process of the dentary is more
angled. A part of these differences supports the hypothesis of even more powerful jaws in A. deinodon than in
the ClB taxon. These two sperm whales were much likely able to feed on other odontocetes, pinnipeds, and
seabirds, using their jaws and full dentition to catch their prey, tear it to pieces, and swallow it.

Key words: sperm whale, Miocene, Peru, jaws, supracranial basin

58

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
IGUANODON'S BRAIN AND PERSPECTIVES ON ORNITHOPOD
EVOLUTION

Pascaline LAUTERS 1, 2, Walter COUDYZER 3, Martine VERCAUTEREN 2, Pascal
GODEFROIT 1

1 Royal Belgian Institute of Natural Sciences, Department of Paleontology, rue Vautier 29, 1000 Brussels,
Belgium, plauters@ulb.ac.be, pascal.godefroit@naturalsciences.be
2 Université Libre de Bruxelles, Laboratoire d'Anthropologie et Génétique humaine, avenue F.D. Roosevelt
50, 1050 Brussels, Belgium, mvercau@ulb.ac.be
3 Universitaire Ziekenhuis Gasthuisberg, Radiologie, Herestraat 49, 3000 Leuven, Belgium,
cwscan@yahoo.com

The brain is generally lost little time after the death of the animal. In particular conditions, the brain or a
natural endocranial cast can be preserved and be fossilized1, 2. Only few insights of the brain of Iguanodon were
available previously3, 4, 5. Unfortunately no detailed descriptions were done on these specimens and the brain
morphology of this dinosaur is poorly understood. CT scan is a medical technology that allows accessing the
internal parts of the fossils, especially the complex braincase of dinosaurs, without damaging them. It is therefore
now possible to investigate the complex braincase of the dinosaurs and to reconstruct the brain and the inner ear
of these extinct animals. Digital endocasts increase the number of specimens studied. Since 2005, CT scan is
used at the Royal Belgian Institute of Natural Sciences in order to reconstruct the evolution of the brain of
Cretaceous ornithopod dinosaurs, among which the collection of Bernissart's Iguanodons. Analysis of the brain
morphology brings new insight about the evolution of the brain and the interrelationships of the major
ornithopod lineages.

1. KUROCHKIN, E.V., SAVELIEV, S.V., POSTNOV, A.A., PERVUSHOV, E.M., & POPOV, E.V., 2006. On the brain of a primitive bird
from the Upper Cretaceous of European Russia. Paleontological Journal, 40(6): 655-667.
2. ROGERS, S.W., 1998. Exploring dinosaur neuropaleobiology: viewpoint computed tomography scanning and analysis of an Allosaurus
fragilis endocast. Neuron, 21: 673-679.
3. NORMAN, D.B., 2005. Dinosaurs: a very short introduction. Oxford University Press, Oxford, 176 p.
4. HULKE, J.W., 1871. Note on a large reptilian skull from Brooke, Isle of Wight, probably dinosaurian, and referable to the genus
Iguanodon. Quarterly Journal of the Geological Society, 27: 199-206.
5. ANDREWS, C.W., 1897. Note on a cast of the brain-cavity of Iguanodon. The Annals and Magazine of Natural History, 114: 585-591.

Key words: Iguanodon, Ornithopoda, Cretaceous, brain, CT scan

59

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
DIAGENESIS OF THE FOSSIL BONES OF IGUANODON
BERNISSARTENSIS

Thierry LEDUC

Royal Belgian Institute of Natural Sciences, Vautier street 29, B-1000 Bruxelles, and Laboratory of
Mineralogy B.18, University of Liège, B-4000 Liège, Belgium, thierry.leduc@sciencesnaturelles.be

Bone diagenesis is the alteration of the chemical composition of the skeleton after the burial.
Specifically, diagenesis is the result of cumulative physical, chemical and biological processes that will modify
the original chemical and/or structural properties of an organic object and will govern its ultimate fate, in terms
of preservation or destruction.
Bone fossilisation is a complex process that includes the degradation of organic matter, the
recrystallisation of bone apatite, the enrichment in trace-elements, the precipitation of new minerals in bone
cavities and finally the compaction process.
The fossil bones of Iguanodon bernissartensis, buried in shale about 125 million years ago, were
discovered in a gallery of the Bernissart mine in 1878. Thirty complete specimens were collected, and some of
them are exposed in their natural position at the Royal Belgian Institute of Natural Sciences, while others are in
"burial position" (their original position in the mine gallery). Since their discovery, the fossil bones were treated
in several ways in order to keep their cohesion and to reduce oxidation by air contact or by variation of relative
humidity. The treatments however didn't prevent the alteration of pyrite and the formation of secondary sulphate
minerals.
Pyrite was very abundant in iguanodon fossil bones and made the bones very brittle. The curettage of
the visible pyrite didn't prevent the oxidation of the rest of it and this still increased further the fragility of the
bones. Pyrite oxidation can be limited to iridescence on its surface but it can also transform pyrite into sulphate
minerals. The growth of those sulphates can completely break the bones.
About fifty samples have been studied. The investigation were made with powder X-ray diffraction
techniques (Philips PW-3710 diffractometer, Debye-Scherrer camera and Gandolfi camera, FeK = 1.9373 Å),
scanning electron microscopy, energy dispersive spectrometry and infrared spectroscopy which allowed us to
identify 13 oxidation products of pyrite: more particularly the sulphate minerals szomolnokite (FeSO4.H2O),
rozenite
(FeSO4.4H2O), roëmerite (Fe2+Fe3+2SO4.14H2O), jarosite (KFe3(SO4)2(OH)6), natrojarosite
(NaFe3(SO4)2(OH)6), halotrichite (Fe2+Al2(SO4)4.22H2O), tschermigite [(NH4)Al(SO4)2.12H2O)], melanterite
(FeSO4.7H2O), coquimbite (Fe3+2(SO4)3.9H2O), metavoltine [(K,Na)8Fe2+Fe3+6(SO4)12O2.18H2O], gypsum
(CaSO4.2H2O), anhydrite (CaSO4), and also sulphur (S).
Szomolnokite and rozenite are the most abundant of these minerals and occur in nearly all samples.
These two minerals are only different by the degree of hydration and can transform into each other. This
mechanism is dependant of the relative moisture of the environment.
We also identified other minerals like carbonate-fluorapatite [Ca5(PO4,CO3)3(F,OH)] (recrystallization
of the bone carbonate-hydroxyapatite), sphalerite (ZnS), hemimorphite (Zn4Si2O7(OH)2.H2O), barite (BaSO4),
and quartz (SiO2). The last one is already present in all the samples. The presence of quartz is the result of both
the filling of bone voids and cracks by shale and the precipitation of authigenic crystals and grains.
Pyrite and barite seemed to form in bone cavities when the iguanodons were buried in shale, i.e. in an
anaerobic environment. Sometimes, in a limited area (1 cm3) of the bone, some cavities are empty while others
are full or partially full of pyrite or barite or both. In the last case, pyrite seemed to be formed first.
Apatite that formed during the bone recrystallization is carbonate-fluorapatite; a small amount of CO 2-
3
has been detected by infrared spectroscopy.
By now, we can state that about twenty minerals have been identified in iguanodon fossil bones by
using these methods. Some of them formed in the bone during the burying process while sulphates and other
secondary minerals appeared after the discovery of the iguanodons, as a result of pyrite oxidation. Their
localisation will be investigated in polished and thin sections.

Key words: Iguanodon, diagenesis, pyrite, oxidation, sulphate minerals

60

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
INSULARITY AND DWARFISM IN LATE CRETACEOUS
EUROPEAN DINOSAURS

Jean LE LOEUFF

Musée des dinosaures, 11260 Espéraza, France, jeanleloeuff@yahoo.fr

Late Cretaceous European dinosaurs lived on an archipelago consisting of many islands with a
fluctuating geography as these islands were separated by shallow epicontinental seaways. Following pioneer
work by Nopcsa, many researchers still consider that the whole dinosaurian fauna was affected by insular
dwarfism. Recent excavations in Southern France and Spain however have yielded very large saltasaurid and
rhabdodontid bones or teeth. Campanian and Maastrichtian localities in Provence and Languedoc often yield
middle-size dinosaurs with an average length close to one meter for saltasaurid femora. These bones belonged to
10 to 12 meters long dinosaurs, which is relatively small by sauropod standards. Many isolated bones and at least
two partial skeletons from the Upper Aude Valley and the Aix-en-Provence area are much larger and show that
European saltasaurids could reach 20 meters in length. This is in agreement with size estimates calculated from
the ichnological data of the Fumanya tracksite in Northern Spain where more than 3 000 sauropod tracks have
been recognized. Improved sampling thus does not confirm that latest Cretaceous European dinosaurs were
consistently smaller than the Early Cretaceous dinosaurs and does not support hypotheses of a global dwarfism
among Late Cretaceous European dinosaurs.

Key words: sauropods, Late Cretaceous, Europe, insularity, dwarfism

61

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
MISSISSIPIAN FORMATIONS AND CRETACEOUS TECTONICS: THE
IMPACT OF THE CURRENT GEOTHERMAL RESERVOIR GENESIS
ON THE PAST LANDSCAPES OF THE MONS BASIN

Luciane LICOUR

Faculté Polytechnique de Mons, rue de Houdain, 9, 7000 Mons, luciane.licour@fpms.ac.be

The well-known Bernissart Iguanodons were discovered during mining operations in the Hainaut coal
basin, in the filling of a so-called "natural pit". This natural pit is one of the numerous sinkholes that miners
encountered as the coal exploitation was active, and that are described and localized in the Delmer and Van
Wichelen inventory (Delmer & Van Wichelen, 1980). The natural pits are irregular cylindrical and roughly
vertical structures, with a diameter ranging from several meters to hundreds of meters for the greatest ones. The
brecciated Pennsylvanian sediments they contain is often sealed by Early Cretaceous deposits, i.e. Wealden
facies and Albian "meule", that indicate the period when the pits reached the surface.

The pit occurrence matches with the Palaeozoïc basement lows. As showed by Stevens & Marlière
(1944), the basement appears as a large irregular basin, underlined by the natural pits. Deep dissolution in the
Mississipian underlying soluble rocks are one of the several processes that are thought to play a major role in
regional subsidence.

The Mississipian deposits crop out at the North of the Mons Basin. They show mainly carbonated
series, and a few stratified breccia layers located in the Middle and Late Visean. The Saint-Ghislain borehole,
described by Groessens et al. (1979), gave precious informations about Visean series, as it revealed the presence
of thick anhydrite layers between 1.900 and 2.500 m of depth. Highly permeable karstic breccias discovered
under the anhydrites turned this geological exploration borehole into a geothermal well and started the
exploitation of the deep Dinantian reservoir as an energy source.

The origin of the high permeable layer of Saint-Ghislain is of course the main issue in determining the
reservoir properties. This raises the question of deep karstification in the Hainaut Mississipian formations. From
both field investigation on surface karst (Vergari & Quinif, 1997) and natural pit fillings, it appears that the
whole aquifer allowed intense fluid flow mainly during extensional tectonic regime in the Late Jurassic/Early
Cretaceous times. The pits could be the result of quick anhydrite dissolution that initiated at the intersection of
faults or fractures and then propagated using the interface between carbonates and sulfates, causing the whole
region to subside.

At a larger scale, the active area of dissolution in the deep Dinantian rocks moved southward through
geological times. Evidences for this are the dip angle of some natural pits located on the northern edge of the
basin and the occurrence of the wealden facies deposits in the same area. These facts are regarded as relicts of
the early subsidence.

Also interesting is that thick anhydrite layers still remain at Saint-Ghislain, whereas it seems to have
been dissolved away in both northern and southern adjacent areas. Tectonic influence may explain both the
abnormal thickness of the anhydrites and their intense deformation. Variscan orientations were recognized by,
amongst others, Rouchy et al. (1984). The above mentioned distribution of natural pits south of the Saint-
Ghislain area can thus be related to the same tectonic disorder.

STEVENS C. & MARLIÈRE R., 1944 Révision de la carte du relief du socle paléozoïque du Bassin de Mons. ASGB, T. LXVII, pp 145-
175.
GROESSENS E., CONIL R., HENNEBERT, M., 1979 Le Dinantien du Sondage de Saint-Ghislain. Stratigraphie et Paléontologie. Mem.
Expl. Cartes Géol. Min. Belg., n° 22, 137 p.
DELMER A. & VAN WICHELEN P., 1980 Répertoire des puits naturels connus en terrains houiller du Hainaut. SGB, PP n°172, 79 p.
ROUCHY J.-M., GROESSENS E., LAUMONDAIS A., 1984 Sédimentologie de la formation anhydritique viséenne de Saint-Ghislain
(Hainaut, Belgique). Implications paléogéographiques et structurales. BSBG, T.93, fasc. 1-2, pp 105-145.
VERGARI A., QUINIF Y., 1997 - Les paléokarsts du Hainaut. Geodinamica Acta. 10, 4, pp 175-187.

Key words: natural pits, Mons Basin, Saint-Ghislain borehole, Dinantian anhydrites, subsidence

62

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
REARING FOR FOOD? KINETIC/DYNAMIC MODELING OF
BIPEDAL/TRIPODAL POSES IN SAUROPOD DINOSAURS

Heinrich MALLISON

Museum für Naturkunde Leibnitz-Institut für Biodiversitäts- und Evolutionsforschung an der Humboldt-
Universität zu Berlin, Invalidenstrasse 43, 10115 Berlin, Germany, heinrich.mallison@museum.hu-berlin.de

Popular art and movies often portray sauropods as able to rear into a bipedal or tripodal (with tail support)
pose for feeding or inter- and intraspecific combat. Such suggestions can also be found in the scientific literature
(e.g. Borsuk-Bialynicka, 1977; Bakker, 1986), and Alexander (1985) attempted to estimate rearing ability in
diplodocids based on the position of their Center of Mass (CoM). Rearing would give sauropod an enormous
height range from which to select their food, although the long necks already give quadrupedal sauropods great
reach. Although Stevens & Parrish (2005) argued that sauropod necks were fairly stiff, Christian & Dzemski
(2007) concluded that their mobility was high. In either case, a bipedal pose would have greatly increased the
maximum feeding height for non-brachiosaurid sauropods and open access to food sources beyond the reach of
other animals.
To date, no investigation of the biomechanics of sauropod rearing has been published. I used
kinetic/dynamic computer modeling to assess the mechanics of attaining a bipedal pose, and the torques required
to sustain them for a prolonged time in Diplodocus and Brachiosaurus. The two genera form the extremes of the
wide range of sauropod bauplans, with short forelimbs in Diplodocus and the longest known forelimbs of all
sauropods in Brachiosaurus. The retention of an upright pose was investigated using a quasi-static approach, and
compared to extent elephants.
Several osteological characters of titanosauriforms, some of which are also present in brachiosaurids, are
supposedly connected to rearing ability: flared ilia, a wide range of motion of the femur in the acetabulum, and
opisthocoelus anterior caudals as well as a fused pubic symphysis (Borsuk-Bialynicka, 1977). Wilson & Carrano
(1999) mentioned the wide gauge of titanosaurs as helpful for rearing. The modeling results confirm that a wide
gauge indeed increases stability, but suggest that none of the others is connected to rearing. To the contrary,
higher mobility in the post-cervical vertebral column may be detrimental, because it makes higher muscle forces
in the axial musculature necessary. Aside from the extremely posterior position of the CoM, Diplodocus shows
further adaptations that combine with upright feeding: the most mobile neck in Sauropoda, a proportionally large
pelvic girdle for large muscles, and chevrons with a highly derived shape that allows supporting weight on them
in that area of the tail that touches the ground when the animal is in an upright pose.
Modeling of the rearing motion showed that Brachiosaurus would have had serious stability problems,
due to the large moment arm of its CoM around the hip joint. In Diplodocus the CoM is located close to the hips,
and slight imbalances do not result in large accelerations that threaten to topple the animal. The modeling results
also indicate that diplodocids could sustain a bipedal/tripodal pose with significantly less effort than elephants,
while Brachiosaurus would have to produce extreme torques in the limb joints. Sufficient muscle mass to
produce these can not be placed on the bones, even assuming that dinosaurs were much more muscular than
mammals. It therefore appears likely that diplodocids were the only sauropods capable of prolonged upright
feeding.

ALEXANDER, R. M., 1985. Mechanics of posture and gait of some large dinosaurs. Zoological Journal of the Linnean Society, 83: 1-25
BAKKER, R. T. 1986. The Dinosaur Heresis. William Morrow and Company, New York.
BORSUK-BIALYNICKA, M., 1977. A new camarasaurid sauropod Opisthocoelicaudia skarzynskii, gen.n.sp.n.from the Upper Cretaceous
of Mongolia. Palaeontolica Polonica, 37: 5-64.
CHRISTIAN, A. & DZEMSKI, G., 2007. Reconstruction of the cervical skeleton posture of Brachiosaurus brancai Janensch 1914 by an
analysis of the intervertebral stress along the neck and a comparison with the results of different approaches. Fossil Record, 10:
38-49.
STEVENS, K. A. & PARRISH, J. M., 2005. Neck posture, dentition, and feeding strategies in Jurassic sauropod dinosaurs. In: Carpenter, K.
& Tidwell, V. (eds.), Thunder Lizards: The Sauropodomorph Dinosaurs. Indiana University Press, Bloomington, pp. 212-232.
WILSON, J. A. & CARRANO, M. T., 1999. Titanosaurs and the origin of "wide-gauge" trackways; a biomechanical and systematic
perspective on sauropod locomotion. Paleobiology, 25(2): 252-267

Key words: sauropod, rearing, bipedal, kinetic/dynamic modeling

63

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
A REEVALUATION OF THE EVIDENCE SUPPORTING AN
UNORTHODOX HYPOTHESIS ON THE ORIGIN OF EXTANT
AMPHIBIANS

David MARJANOVI 1, Michel LAURIN 2

1, 2 UMR 7179, Équipe "Squelette des Vertébrés", CNRS/Université Paris 6/MNHN/Collège de France,
4 place Jussieu, case 19, 75005 Paris (France), 1 david.marjanovic@gmx.at, 2 michel.laurin@upmc.fr

The origin of the frogs, salamanders, and caecilians is controversial. McGowan (2002) published an
original hypothesis on lissamphibian origins: Gymnophiona was nested inside the "microsaurian" lepospondyls,
this clade was the sister-group of a caudate-salientian-albanerpetontid clade, and both together were nested
inside the dissorophoid temnospondyls. We have investigated McGowan's data matrix and disagree with the
scoring of 35% of the cells. All taxa and all but two characters are affected. In some cases, we have a different
interpretation about correspondence between morphology and character states, or we delimit states differently
(or use information that was unknown in 2002); furthermore, unlike McGowan, we have decided for each
multistate character separately whether and how to order it, using the stepmatrix gap-weighting method
recommended by Wiens (2001) for the two most obviously continuous characters. In other cases, we report
probable typographic errors. When these cells and characters are revised, the most parsimonious trees (Figure: c)
now longer by almost 64% support one of the three commonly advocated hypotheses, namely a
monophyletic Lissamphibia which is nested, together with its sister-group Albanerpetontidae, within the
temnospondyls (next to Doleserpeton) even though we did not add any characters or taxa to the very small data
matrix. This exemplifies the impact of errors in data matrices on the results of phylogenetic analyses. Adding the
only well-known lysorophian Brachydectes, however, results in the Lissamphibia-Albanerpetontidae clade
becoming the sister-group of Brachydectes and settling within the lepospondyls rather than the temnospondyls,
thus supporting another of the previously published three hypotheses (even though we reinterpret the homology
of some skull bones of Brachydectes as less lissamphibian-like than previously thought). This latter finding does
not change if the recently described Gerobatrachus is also added. Finally, when Doleserpeton is interpreted as
morphologically immature (which means scoring three characters as unknown instead of known), Lissamphibia
and Albanerpetontidae are again nested within the "microsaurian" lepospondyls, even though Brachydectes is
not included in this analysis. This, too, does not change if Gerobatrachus is added and likewise treated as
morphologically immature. Except for Lissamphibia + Albanerpetontidae, Batrachia, Branchiosauridae, Apateon
+ Schoenfelderpeton, Gymnarthridae + Rhynchonkos, and to a lesser degree Lissamphibia, bootstrap supports are
at or below 50% under all assumptions. Such lability had to be expected from the small size of the data matrix.

MCGOWAN GJ. 2002. Albanerpetontid amphibians from the Lower Cretaceous of Spain and Italy: a description and reconsideration of their
systematics. Zoological Journal of the Linnean Society 135: 132.
WIENS JJ. 2001. Character analysis in morphological phylogenetics: problems and solutions. Systematic Biology 50: 689699.

c

Figure: Strict consensus trees resulting from McGowan's original dataset when all characters are unordered (a)
or all multistate characters are ordered according to the original state numbers (b); strict consensus tree resulting
from the modified dataset under the first of five assumptions, namely when Brachydectes and Gerobatrachus are
not added and Doleserpeton is treated as adult and not paedomorphic (c). Extant OTUs in bold.

Key words: Amphibia, Lissamphibia, Brachydectes, Gerobatrachus, stepmatrix gap-weighting

64

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
VEGETATION FROM LATE CRETACEOUS DINOSAUR
ECOSYSTEMS IN AMUR RIVER REGION (RUSSIA AND CHINA)

Valentina MARKEVICH 1, Eugenia BUGDAEVA 1, Yury BOLOTSKY 2

1 Institute of Biology and Soil Science, FEBRAS, 159, Prosp. 100-letiya,Vladivostok, 690022,
bugdaeva@ibss.dvo.ru
2 Institute of Geology and Natural Resources, FEBRAS, 1, Ryolochny Per., Blagoveshchensk, 675000,
dinomus@ascnet.ru

Dinosaur fossils have been found in Santonian and Maastrichtian deposits of the Zeya-Bureya Basin in
Far-Eastern Russia. Dinosaur footprints and bones were discovered in the Santonian Yongancun Formation, and
numerous bones and skin remains were unearthed in the Maastrichtian Yuliangzi Fm (China) and Lower
Tsagayan Subformation (Russia).
During the Santonian the Zeya-Burey Basin was occupied by a vast, but shallow lake. On the swampy
of this lake, cycadophytes, taxodialeans, conifers and cyatheaceous ferns grew. The slopes were covered by
representatives of Pinaceae, Podocarpaceae, Juglandaceae, Fagaceae, Platanaceae and plants producing «unica»-
pollens. Probably, Dicksoniaceae and Osmundaceae were in understory.
The lake reduced during the Campanian, while the relief and the vegetation changed. The frame of
basin increased. The vegetation was formed by conifers and, in understory, ferns Cyatheaceae and
Polypodiaceae, sometimes bryophytes. Angiosperms remained poorly represented, but their diversity became
more important.
The rising of the Zeya-Bureya area continued during the Maastrichtian. River valleys with numerous
small lakes appeared. The vegetation of the first half of this stage was abundant and diverse. Plant communities
of these valleys were represented by bryophytes, ferns (Ophioglossaceae and Cyatheaceae), gymnosperms
(Pinaceae, Taxodiaceae and diverse gnetaleans). The slopes of the valleys were covered by forests, dominated by
angiosperms that mainly produced tricolpate pollen. Plants producing «unica»-type pollen, were rare, but
diversified.
In the `mid'-Maastrichtian this type of vegetation persisted, but the role of angiosperms increased: they
became dominant, sometimes reaching 44% of the vegetation. The taxonomical composition of angiosperms
considerably changed and flowering plants related to families Betulaceae, Juglandaceae, Fagaceae, Ulmaceae,
Myricaceae prevailed.
At the end of the Maastrichtian, the palaeoenvironments changed. The uplift of this area reached its
maximum. The climate became more contrasted, humid and less warm. The vegetation was dominated by
taxodialeans and representatives of Ulmaceae. Among spores monolete Laevigatosporites predominated.
Angiosperms were diversied and represented by taxa close to Betulaceae, Salicaceae, Juglandaceae, Myricaceae,
and Myrtaceae. The diversity of «unica»-type pollens increased, sometimes reaching 33% of the palynological
spectrum.
The Santonian, Campanian and first half of the Maastrichtian stages were favorable for dinosaurs, but
then the fall of temperature and the more contrasted climates provoked the destruction of these ecosystems and
the extinction of the dinosaurs.

Key words: Cretaceous, vegetation, dinosaurs, Zeya-Bureya basin

65

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
3D MODELLING OF THE PALEOZOIC TOP SURFACE IN THE
BERNISSART AREA AND INTEGRATION OF DATA FROM
BOREHOLES DRILLED IN THE "CRAN AUX IGUANODONS"

Thierry MARTIN 1, Johan YANS 2, Christian DUPUIS 1, Paul SPAGNA 1,
Olivier KAUFMANN 1

1 FPMS, GFA, 9 rue de Houdain, 7000 Mons, Belgium, Thierry.Martin@fpms.ac.be,
Christian.Dupuis@fpms.ac.be, Paul.Spagna@fpms.ac.be, Olivier.Kaufmann@fpms.ac.be
2 FUNDP UCL-Namur, Geology, 61 rue de Bruxelles, 5000 Namur, Belgium, johan.yans@fundp.ac.be

Since 1878-1881 and the discovery of numerous complete skeletons of dinosaurs in Bernissart, several
studies have been dedicated to the paleontological content of the "Cran aux Iguanodons" (see references in
Bultynck, 1989). However little is known about the geometry of the Cran and its integration within the Paleozoic
basement of the Mons Basin. In 2002-2003, three new boreholes (BER2, BER3 and BER4) allowed us to
improve our understanding of the geometry of the Cran, integrating the new data in a 3D model of the top
surface of the Paleozoic basement.
In the BER2 core (Belgian Lambert coordinates X=98.910, Y= 130.021, Z=33) the Wealden facies have
been sampled from -265.5 meters of depth. However, diagraphic analysis (Legrain & Tshibangu, 2004) indicate
that the top of the wealden strata in BER2 is localised almost 40 meters higher, around -227 meters. In the same
borehole, Namurian basement has been cut at -291 meters. The BER3 core (Belgian Lambert coordinates:
X=98.870, Y=129.961, Z=24) cuts the Wealden facies at -265 meters of depth and reached the Carboniferous
basement at -315 meters. The BER4 core (Belgian Lambert coordinates X=98.910, Y=129.961, Z=24) did not
reach the natural pit, the clayey sediments found under the Meule facies at -246 m being attributed to the
weathered Namurian basement.
In a first step, the 3D modelling of the top surface of the Paleozoic basement is created at a regional scale
(area of 340km²) from outcrop limits and boreholes descriptions. In this way the overall geometry is outlined.
The studied area is located in the Western part of Mons Basin and near the French border. The methodology is
implemented from a system architecture with file transfers using ArcGIS, Gocad and a database (Kaufmann &
Martin, 2008). Boreholes descriptions are collected, structured and stored in a database. In ArcGIS, boreholes
are positioned and formation limits are drawn from digitized geological maps. In this first step, these data are
imported in Gocad in order to build a surface in a 3D space covering the whole area. In a second step, this model
is refined at local scale (~11 km²) in the Bernissart area around the Cran including data from new boreholes
(BER2, BER3 and BER4).

Figure: 3D model of the top surface of the Paleozoic basement at regional scale with data from the new borehole
BER3.

BULTYNCK, P., 1989. Bernissart and the Iguanodons, Publication of the RBINS, Brussels. 115 p.
LEGRAIN, H. & TSHIBANGU, J.-P., 2004. Sondages exploratoires dans le cran aux iguanodons de Bernissart : Rapport sur les mesures
géophysiques (diagraphie en forage et tomographies sismiques entre trous). Service de Génie Minier, Faculté Polytechnique de Mons,
50 pages + annexes.
KAUFMANN, O. & MARTIN, T., 2008. 3D geological modelling from boreholes, crosssections and geological maps, application over
former natural gas storages in coal mines. Computers & Geosciences, 34: 278-290.

Key words: Bernissart, boreholes, Mons Basin, natural pit, 3D modelling

66

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
STRATIGRAPHY OF THE HAINE GROUP ("MEULE" SEDIMENTS)
OVERLYING THE WEALDEN FACIES IN THE BER 3 BOREHOLE
(BERNISSART, MONS BASIN, BELGIUM)

Edwige MASURE 1, Johan YANS 2

1 Université Pierre- Marie CURIE, UMR-CNRS 5143, 4 pl. Jussieu, 75005 Paris, France,
edwige.masure@upmc.fr
2 FUNDP, UCL-Namur, Rue de Bruxelles 61, 5000 Namur, Belgium, johan.yans@fundp.ac.be

Between 75 and 265 meters of depth in the BER 3 borehole drilled in 2002-2003, the sediments have the
typical facies of the Haine Group, locally called "Meule". In its stratotype, located in the Harchies borehole (
2.9 km far from the BER 3 borehole), the 173.2 m-thick "Meule" is divided into five Formations, dated by their
content of ammonites (Amédro, 2002; from base to top): Formation of the Green sandstones of Pommeroeul
(Middle Albian), Formation of Harchies, Formation of Catillon (both Upper Albian sensu stricto), Formation of
Bracquegnies (Vraconnian) and Formation of the Calcirudites of Bernissart (Early to Middle Cenomanian).
These latter directly cover the dinosaurs-bearing Wealden facies. In the BER 3 borehole, the "Meule" shows the
following main facies (Yans et al., 2005):
(1) 265 - 170.7 m of depth: argillaceous and calcareous glauconiferous sandstone and calcarenite, locally silicified, rich in
sponges. Local cm-voids and fossiliferous beds. From 258,4 m to base: conglomerate. Harchies Formation.
(2) 170.7 - 135.4 m: glauconiferous calcarenite with trigonia, locally indurated. Some green conglomeratic levels (mainly at
the base). Catillon Formation.
(3) 135.4 - 105.3 m: alternances of sandy grey fossiliferous, glauconiferous calcarenite and conglomeratic calcarenite.
Bracquegnies Formation.
(4) 105.3 - 75 m: sandy, glauconiferous calcarenite, locally indurated, cellular and geodic, bioclastic levels. Local flints and
pebbles. Calcirudites of Bernissart Formation.
These facies are similar to those described in the Harchies stratotype (Robaszynski et al., 2001). In the
BER3 borehole, the "Meule" may lack the Formation of the Green sandstones of Pommeroeul. The incomplete
"Meule" of the BER3 is however thicker (190 m) than the one at the Harchies stratotype (173,2 m). These local
thickness variations are due to the different subsidence rates and centres in the Mons Basin during the deposition
of the Wealden facies and Meule sediments (Barremian to middle? Cenomanian), in relation with deep anhydrite
dissolutions and/or extensive regional crustal activity (Spagna et al., 2007).
The age of the "Meule" at the BER3 borehole is determined by its content of dinoflagellate cysts. The
biostratigraphy is based on first appearance data (FAD) of boreal index species which are correlated with the
ammonite and foraminiferal zonal schemes (Foucher & Monteil in Hardenbold et al., 1998). The occurrence of
Surculosphaeridium ?longifurcatum at 261,6 m indicates that the sample can not be older than the Middle
Albian. Litosphaeridium conispinum at 252.6m suggests that this level is in the upper part of the Middle Albian.
In sample 155.2m, the occurrence of Xiphophoridium alatum suggests an Upper Albian age. Litosphaeridium
siphoniphorum is recorded in sample 129.2 m, the FAD of the species is located at the base of the Dispar zone
(Vraconnian). Epelidosphaeridia spinosa is observed in sample 102.7 m. The FAD of the species is known in the
Vraconnian.
On the basis of these preliminary results, we suggest the following biostratigraphy:
(1) 261.6 - 252.6 m: Harchies Formation (or Green sandstones of Pommeroeul Fm?). Upper part of Middle
Albian.
(2) 155.2 - 138.1 m: Catillon Formation. Upper Albian.
(3) 129.2 m: Bracquegnies Formation. Vraconnian.
(4) 102.7 m: Calcirudites of Bernissart Formation. Vraconnian or younger (most probably younger).

AMEDRO, F., 2002. Plaidoyer pour un étage Vraconnien entre l'Albien sensu stricto et le Cénomanien (système Crétacé). Memoir of the
Royal Academy of Belgium. 128 p.
FOUCHER, J.-C. & MONTEIL, E., 1998. Cretaceous biochronostratigraphy, Chart 5; Mesozoic and Cenozoic Sequence Chronostratigraphic
Framework of European Basins. In: de Graciansky, P.-C., Hardenbol, J., Jacquin, T., & Vail, P.-R. (Eds.), Mesozoic and Cenozoic
Sequence Stratigraphy of European Basins, SEPM Special Publication: 60.
SPAGNA, P., VANDYCKE, S., YANS, J. & DUPUIS, C. 2007. Hydraulic and brittle extensional faulting in the Wealden facies of
Hautrage (Mons Basin, Belgium). Geologica Belgica, 10/3-4: 158-161.
ROBASZYNSKI, F., DHONDT, A., & JAGT, J.W.M., 2001. Cretaceous lithostratigraphic units (Belgium). In: Bultynck, P. & Dejonghe, L.
(eds), Guide to a revised lithostratigraphic scale of Belgium, Bruxelles, Geologica Belgica: 121-134.
YANS, J. et al. 2005. Description et implications géologiques préliminaires d'un forage carotté dans le "Cran aux Iguanodons" de Bernissart.
Geologica Belgica, 8 : 43-49.

Key words: Dinoflagellates, Bernissart, Mons Basin, Belgium, Haine Group

67

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
COMPOSITION AND BIOGEOGRAPHIC AFFINITIES OF THE
MIDDLE EOCENE MESSEL AVIFAUNA

Gerald MAYR

Forschungsinstitut Senckenberg, Sektion Ornithologie, Senckenberganlage 25, D-60325 Frankfurt/M.,
Germany, Gerald.Mayr@senckenberg.de

Birds are among the most abundant land vertebrates in the Middle Eocene fossil site Messel in
Germany, and so far more than 500 skeletons have been found, which belong to over 50 species. The talk gives
an overview of the composition of the Messel avifauna and its biogeographic affinities. It has been known since
a while that compared to coeval avifaunas the Messel avifauna closely resembles that of North America, but only
recently the striking degree of congruence has been appreciated. Many of the shared arboreal taxa disappeared in
North America towards the Oligocene, whereas they persisted in the Old World. A proportionally high number
of avian taxa from Messel has its closest extant relatives in the Southern Hemisphere, especially South America.
Disappearance of the stem group representatives of these lineages from the Northern Hemisphere cannot be
explained by climatic cooling during the Cenozoic alone, and a better knowledge of Paleogene avifaunas will
certainly lead to an increased understanding of the evolution of Cenozoic ecosystems.

Key words: Messel, birds, Eocene, biogeography

68

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE BERNISSARTIDS: COMMON EUROPEAN CRUSHING
CROCODILES

Jean-Michel MAZIN 1, Joane POUECH 1, Julien RASLAN-LOUBATIE 2

1 UMR 5125 CNRS, Université Lyon1, Campus de la Doua,
Bâtiment Géode, 69622 Villeurbanne cedex, France, jean-michel.mazin@univ-lyon1.fr,
joane.pouech@pepsmail.univ-lyon1.fr
2 17, rue Raspail, 92300 Levallois, France, loubatie@gmail.com

The site of Cherves-de-Cognac (Berriasian, Charente, France) is characterized by its rich vertebrate
fauna among which crocodylomorphes are abundant. Four families are present: Atoposauridae, Bernissartidae,
Pholidosauridae and Goniopholididae, from macroremains as well as from microremains.
Among the numerous crocodylomorph teeth found at Cherves-de-Cognac, many can be referred to the
tribodont morphotype classically considered as bernissartid posterior teeth. However, numerous more or less
elevated conical teeth showing the same enamel ornamentation are associated with these rounded crowns and
might represent anterior and intermediate bernissartid teeth. As already noted by Buffetaut & Ford (1979) and
described by Brinkmann (1992), three morphotypes can be distinguished among the Bernissartia-like teeth:
anterior conical teeth, intermediate leaf-shaped teeth and posterior bulbous tribodont teeth.
Bernissartids are very common at Cherves-de-Cognac, since 5360 isolated teeth have been found from
28 levels of the section. This great number of specimens, showing the variation of crown morphology along the
tooth row, shows teeth at different stages of wear, from unworn bulbous teeth with carinae to strongly worn
crowns.
Such particular isolated bulbous teeth have been reported from several other Late Jurassic to Early
Cretaceous European sites, namely: Guimarota (Kimmeridgian, Portugal), Chassiron (Lower Tithonian, France),
Boulonnais (Upper Tithonian, France), Galve (Upper Tithonian, Spain), Cherves-de-Cognac (Lower to Middle
Berriasian, France), Middle Purbeck Limestone Group (Lower and Middle Berriasian, England), Unã
(Barremian, Spain), Isle of Wight (Barremian, England), Bernissart (Upper Barremian-Lower Aptian, Belgium),
Vallipón (Upper Barremian, Spain) and Buenache de la Sierra (Upper Barremian, Spain). These isolated
tribodont teeth are always referred to the genus Bernissartia or to the family Bernissartidae. However, Salisbury
(2002) questions the generic identification of these rounded teeth, and notably the specimens from Sunnydown
Sly Bed of Middle Purbeck Limestone Group. He argues that this attribution is only based on the similitude of
the enamel ornamentation and apical wear, with the Belgian lectotype. He also notes that several upper
Cretaceous European and North American genera such as Brachychampsa, Albertochampsa, Stangerochampsa
and Allognathosuchus show such posterior tribodont teeth.
However, as far as it is known, the oldest Eusuchian crocodylomorphes are from the Albian-
Cenomanian, and there are some noticeable morphological differences between alligatorid and bernissartid teeth.
For example, intermediate leaf-shaped teeth are a bernissartid feature when they do not exist in alligatorids;
enamel ornamentation, made of numerous strong vertical wrinkles in bernissartid is weakly marked in
alligatorids; posterior rounded teeth are laterally compressed or kindney-shaped in bernissartids, when they are
clearly hemispheric in alligatorids. Thus, they is no doubt that these bulbous isolated teeth found in the Late
Jurassic and Lower Cretaceous of Europe can be referred to the family Bernissartidae, as well as the associated
leaf-shaped and conical morphotypes.
So, the bernissartid crocodyliformes were widely distributed in Western Europe during the Late Jurassic
and the Early Cretaceous in continental to margino-littoral depositions. They were characterized by a particular
heterodont dentition, with crushing bulbous posterior teeth reflecting a diet probably based on hard-shelled
preys, such as freshwater molluscs, crustaceans or strongly scaly fishes such as semionotid, commonly found in
the same faunal assemblages.

BRINKMANN W., 1992. Die Krokodilier-Fauna aus der Unter-Kreide (Ober-Barremium) von Uña (Provinz Cuenca, Spanien). Berliner
Geowissenschaftliche Abhandlungen, E(5), 121 pp.
BUFFETAUT E. & FORD R.L.E., 1979. The crocodilian Bernissartia in the Wealden of the Isle of Wight. Palaeontology, 22: 905-912.
SALISBURY S., 2002. Crocodilians from the Lower Cretaceous (Berriaisian) Purbeck Limestone Group of Dorset, Southern Englanc.
Special Papers in Palaeontology, 68, 121-144.

Key words: Bernissartidae, Late Jurassic, Early Cretaceous, heterodonty, western Europe

69

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE BASAL RADIATION OF RUMINANTIA IN ASIA AND THE
"DICHOBUNOID GORDIAN KNOT"

Grégoire MÉTAIS 1, Tao QI 2, Jianwei GUO 2, Chris BEARD 3

1 Paléobiodiversité et Paléoenvironnements, UMR 5143 (CNRS, MNHN, UPMC), Département Histoire de la
Terre, Muséum National d'Histoire Naturelle, 8 rue Buffon, CP 38, 75231 Paris, France, metais@mnhn.fr
2 Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, P. O. Box 643,
Beijing 100044, China
3 Section of Vertebrate Paleontology, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh,
PA 15213, USA, beardc@CarnegieMNH.org

At present, there is little or no consensus regarding the phylogenetic relationships among primitive
ruminant artiodactyls. Phylogenetically significant morphological characters are thought to be distributed among
the skull and dentition, as well as in limb structure. Incompletely known fossil taxa may, therefore, be especially
problematic for purposes of phylogeny reconstruction. Added to this difficulty is a substantial and well-
documented level of homoplasy within selenodont artiodactyls.
Available molecular evidence supports a relatively ancient dichotomy between living tragulids and
pecorans (horned ruminants). For example, the genomes of tragulids and nonruminant artiodactyls (suborders
`Suiformes' and Tylopoda) lack a short interspersed repeat that is present in all pecoran ruminants examined.
Likewise the phylogenetic relationships between Ruminantia and the other groups of cetartiodactyls remain
unclear from both molecular and paleontological perspectives, and in all cases sources of discrepancies. It
appears that these discrepancies between morphological and molecular data sets are probably biased by the
inconsistence of the current fossil record of artiodactyls in Asia, the region where they probably originated, and
the resulting poor resolution of the phylogenetic relationships between Asian, European, and North American
"dichobunoids", a vast, and loosely delimited group of paraphyletic cetartiodactyls.
One key for resolving discrepancies between morphological and molecular data is to understand the basal
radiation and phylogeny of Selenodontia that appear suddenly in the fossil record during the middle Eocene.
Ruminants derive from a still non-identified group of Asian dichobunoids. It is generally admitted that
dichobunoids represent a heterogeneous and paraphyletic assemblage of generalized bunodont to bunoselenodont
artiodactyls that inhabited the three Holarctic continents during the Paleogene.
Although phylogenetic relationships among the various dichobunoids remain obscure, attempts to resolve
this Gordian knot have been hampered in part by the comparatively sparse Eocene fossil record of Asia. In this
context, the discovery of new taxa of Asian dichobunoids represents a potential advance in understanding the
early phylogeny and biogeography of the group, and in turn the basal radiation of Ruminantia.

Key words: cetartiodactyls, Ruminantia, Eocene, Asia

70

Darwin-Bernissart meeting, Brussels, February 9-13, 2009

CARBONIFEROUS AMPHIBIANS AND REPTILES 
EXPERIMENTATION AND DIVERSIFICATION

Andrew R. MILNER

Department of Palaeontology, The Natural History Museum, Cromwell Road, LONDON SW7 5BD, UK,
andrew.milner@nhm.ac.uk

Following the evolution of terrestriality in tetrapods in the late Devonian and basal Carboniferous (370-
350 Ma), the group diversified during the Carboniferous. Forty million years later, in the late Moscovian stage of
the Carboniferous (310 Ma), there were at least 30 lineages of tetrapod, all filling different niches. Using the
skeletons of early tetrapods and the circumstances in which they occur, we can deduce some of the biological
and ecological basis of this early diversity. In this review, several aspects of this diversification will be outlined.
1. Pattern of respiration. Two mechanisms of respiration can be deduced from the skeleton, namely buccal
pumping and costal breathing. Costal breathing permits the development of a neck and more complex food
processing in the mouth.
2. Size. The earliest tetrapods were about a metre in length. By the late Carboniferous, there were 4 m
long forms and 12 cm long forms. The latter show evidence of progenetic dwarfing in the reduction in the
numbers of skull bones.
3. Degree of terrestriality. Different associations of Tetrapoda are found in different preservational
conditions and terrestrial, marginal/amphibious and aquatic associations can be recognised. Morphological
evidence for lifestyle includes degree of ossification of wrist and ankle bones, presence or absence of lateral-line
canals.
4. Dentition and feeding. In the Moscovian, all tetrapods appear to have been carnivores feeding on
invertebrates or other vertebrates. A range of dental types include piercing, gripping and crushing teeth. The
skulls of early reptiles show evidence of different arrangements of jaw muscles for different feeding techniques.
Immediately after the Moscovian, the first two lineages of unambiguous herbivore appear the edaphosaurs and
the diadectids. These were large-bodied forms with leaf-stripping and crushing dentitions. They probably
digested cellulose by hind-gut fermentation.
5. Pattern of locomotion. Most forms had long undulating bodies and short limbs and would have
progressed using the body for propulsion and the limbs as anchors. A few lineages developed longer limbs and
shorter bodies and would have moved by limb-based thrusting. The aïstopods were limbless analogues of snakes
and had also evolved a snake-like skull.
6. Reproduction and metamorphosis. There is little direct evidence of patterns of reproduction. We know
that frog-like eggs were produced by some amphibians of unknown type. Most early tetrapods show simple
growth and development patterns and only one group of amphibians the Dissorophoidea shows evidence of
a metamorphosis during development. These forms were able to occupy ephemeral niches as larvae.
7. Mechanism for hearing air-borne sound. All early tetrapods could hear low-frequency ground-borne or
water-borne sound. One group the temnospondyls had frog-like ears capable of picking up high frequency
air-borne sound, permitting insect detection and with the potential for producing and hearing calls inaudible to
other tetrapods.
8. Anti-predator defence. One amphibian Stegops shows the development of spines protecting the neck
the earliest known example of an anti-predator defensive structure.

Our understanding of the relationships and phylogeny of these early tetrapods is currently in a state of
flux. Over the next decade we can anticipate that it will stabilise and we will then be able to map these
adaptations onto a `family tree' and gain understanding of the evolutionary biology of tetrapod diversification on
land.

Key words: coal-swamp, evolution, morphology, Moscovian, tetrapods

71

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
NEW DATA ON THE VALANGINIAN-HAUTERIVIAN REPTILE
OOTAXA OF THE IBERIAN RANGE (NE OF SPAIN)

Miguel MORENO-AZANZA 1, José Manuel GASCA 1, José Ignacio CANUDO 1

1 Grupo Aragosaurus. Universidad de Zaragoza. Pedro Cerbuna 12, 50009 Zaragoza, Spain,
mmazanza@unizar.es

Despite the great amount of work on fossil eggshell fragments that has been carried out during the last
two decades, Lower Cretaceous reptile ootaxa are still poorly known, especially in Europe. This is due to both
the lack of well-preserved nests and eggs from this age in the fossil record and the fact that fragmentary material
is not as abundant as in Upper Cretaceous localities. As the prospecting and sieving campaigns undertaken by
our research group Aragosaurus during the last fifteen years have allowed us to accumulate a worth the effort to
study material, the Iberian Range, in NE Spain, has emerged as a priviliged area for filling this gap.

The study area is a small outcrop of the Villanueva de Huerva Formation in a small gully located 2 km
south of the town of Villanueva de Huerva (Zaragoza province) and less than 100 m away from the stratotype
section of this Formation. The Pochancalo 1 locality is a 1-meter-thick bed of grey clays deposited in a fluvio-
lacustrine environment. Fossil remains are relatively abundant and are mainly microvertebrate and plant
fragments. The vertebrate remains identified are fish scales and teeth (semionotiformes and hybodontids),
postcranial amphibian bones, chelonian plates, and crocodilian (Atoposauridae, Bernissartidae and
Goniopholidae families), pterosaur, mammal (multituberculate) and dinosaur (sauropod and theropod) teeth.

Over 200 hundred fossil eggshell fragments have been collected at the Pochancalo 1 locality by sieving
250 kg of sediment. We ascribe most of them to the oofamilies Spheroolithidae, Megaloolithidae,
Prismatoolithidae and Testudoolithidae. Spheroolithidae eggshells have been related to hadrosaurian dinosaurs.
The absence of fossil bones of this group in the Lowermost Cretaceous of Europe suggests that more basal
ornithopods laid Spheroolithid eggshells too.

Noticeable is that eggshells ascribed to Megaloolithidae are profuse in Upper Cretaceous beds, though
also known in the Jurassic of France and Portugal and the Barremian of Spain they are unknown at the basalmost
Cretaceous. The Pochancalo 1 megaloolithid present an intermediate set of characters between Upper Cretaceous
and Jurassic ootaxa. Testudoolithidae eggshells differ from the ootaxa previously known in the Lower
Cretaceous and may also correspond to a new oospecies of Testudoolithus.

Prismatoolithidae eggshells differ from the ootaxa previously known from the literature by their
external ornamentation. They sligthly resemble other eggshells that have been collected in younger localities of
the Iberian Range, uppermost Hauterivian-Lower Barremian in age, and may be part of a new oogenus yet to be
described.

Key words: eggshell, ootaxa, Pochancalo, Lower Cretaceous, Iberian Range

72

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
PALAEOENVIRONMENTAL CONTEXT OF THE `BLACK MARBLE'
OF DENÉE (VISÉAN, BELGIUM)

Bernard MOTTEQUIN 1, Edouard POTY 2

1-2 Unité de Paléontologie animale et humaine, Université de Liège, Bât. B18, Allée du 6 août, Sart Tilman,
B4000 Liège 1, bmottequin@ulg.ac.be and E.Poty@ulg.ac.be

Fish remains such as isolated teeth or scales are uncommon in Tournaisian and Viséan (Lower
Carboniferous) rocks of southern Belgium. Moreover, large fragments or almost complete fishes are particularly
rare and only known in two Viséan localities: Denée (Namur province) and Visé-Richelle (Liège province).
The quarries located around the village of Denée have yielded rare but remarkably preserved fishes and
invertebrates (including echinoderms and dendroid graptolites). The latter have been collected within the `black
marble' of Denée when this black coloured limestone was intensively and manually quarried at the end of the
19th century and at the beginning of the 20th century. If it had not been worked, the `black marble' of Denée
would have been considered probably as azoic due to the rarity of the fossils. As most of the excavations were
subterranean and as they are now flooded and disused, it is not possible to sample all the fauna anymore. Four
fish species have been described for the first time on the basis of material collected in Denée: the chondrostean
Benedenius deneensis and three elasmobranchs (Denaea fournieri, Cratoselache pruvosti, Sphenacanthus
delepinei).
The `black marble' of Denée is now included in the Molignée Formation of Lower Viséan age
(Moliniacian). This formation (c. 60 m thick) consists of a succession of thin-bedded, commonly laminated black
limestones (`black marble') which alternate with thick-bedded, dark-grey limestones (`thick beds'). It developed
in a confined intra-platform basin [central part of the Dinant sedimentation area (DSA)] progressively filled by
distal calcareous turbidites originating from the southward prograding shelf to the north. This basin was bordered
to the south by a discontinuous barrier of Waulsortian mudmounds built against a major synsedimentary fault
separating the DSA from the southern Avesnois sedimentation area. The alternations of laminated and
bioturbated lithofacies occurring within the Molignée Formation implies that the palaeoenvironment recorded
several anoxic to dysoxic periods alternating with more oxygenated ones due to sea-level fluctuations of low
magnitude. This periodic confinement of the central part of the DSA took place during a third-order sequence
characterized by a low sea level, namely the sequence 5 of Hance et al. (2001). Low oxygen concentrations are
also suggested by the existence of dysaerobic organisms such as the bivalves of the `paper pecten' morphotype
and the remarkable preservation of the fauna.
The `black marble' of Denée is a fossil conservation deposit and belongs more particularly to the
`obrution deposits' of Seilacher et al. (1985) (Mottequin 2008). The turbiditic sedimentation with smothering
effect (rapid burial) combined with deficient oxygenation of the bottom waters favoured the exceptional
preservation of the faunas (e.g. fishes, echinoids, ophiuroids) by inhibiting the development of the necrophagous
and saprophagous organisms during the deposition of the `black marble' facies sensu stricto.

HANCE L., POTY E. & DEVUYST F.X. 2001. Stratigraphie séquentielle du Dinantien type (Belgique) et corrélation avec le Nord de la
France (Boulonnais, Avesnois). Bulletin de la Société géologique de France 172: 411426.
MOTTEQUIN B. 2008. The `black marble' of Denée, a fossil conservation deposit from the Lower Carboniferous (Viséan) of southern
Belgium. Geological Journal 43: 197208.
SEILACHER A., REIF W.E. & WESTPHAL F. 1985. Sedimentological, ecological and temporal patterns of fossil-Lagerstätten. In
Whittington H. & Conway Morris S. (eds), Extraordinary Biotas: Their Ecological and Evolutionary Significance. Philosophical
Transactions of the Royal Society of London 311: 523.

Key words: `black marble' of Denée, Carboniferous, Viséan, Belgium

73

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
IGUANODONTIANS FROM THE WEALDEN OF BRITAIN AND
EUROPE

David B. NORMAN

Department of Earth Sciences, University of Cambridge CB2 3EQ, UK

Basal (non-hadrosaurid) iguanodontians are, without doubt, the most important group of ornithischian
dinosaurs to have been discovered, collected and studied in Wealden-aged beds across western Europe. Their
remains were first recognised and described scientifically in 1825 on the basis of disarticulated and dissociated
remains (primarily distinctive teeth) collected from outcrops of "Tilgate Grit" (Horsham Stone: Hauterivian
Stage) in Sussex by Dr and Mrs Mantell (aided by paid labourers working in several quarries in the vicinity of
Whiteman's Green, Cuckfield). While such remains were unique and distinctive at the time, and merited the
creation of a new genus Iguanodon (Mantell, 1825), posterity has demonstrated that teeth of this general
morphology are common to a range of ornithopod ornithischian dinosaurs that have now been identified from
rocks that span much of the Late Jurassic and Early Cretaceous; as well as such geographically disparate regions
as Eurasia, North America and North Africa (Norman, 2004).
Attempts to rationalize the taxonomy of European Wealden-aged ornithopods (and approximate
contemporaries from other geographic regions) has been the long-term aim of a series of studies that have been
produced across the past two decades (Norman, 1980, 1986, 1991, 2002, and references therein). Such studies
necessarily encompass the range of variation exhibited by `osteological species' (consistent operational
taxonomic units) and need to be based on detailed anatomical comparison of known material. Early work
determined that it was appropriate (to ensure historical stability) to preserve the generic name by establishing a
holotype Iguanodon anglicus (Holl, 1829) by determining a lectotype and paratype series from the specimens
described collectively by Gideon Mantell. Subsequently, further material was collected from both lower
exposures (Wadhurst Clay Valanginian Stage) of the Wealden, particularly in the Hastings area (this was
described piecemeal as "Iguanodon mantelli" by Richard Owen in the Palaeontographical Monographs series);
and at the same time occasional remains from the Wealden Marls (Barremian) of the Isle of Wight were also
described by Owen under the same taxonomic name.
The spectacular discovery of fully-articulated ornithopod dinosaurs at Bernissart in the late 1870s
(described in numerous articles by Louis Dollo [1882, et seq] summarised in Norman, 1980, 1986) added
enormously to the general understanding of the anatomy of these types of dinosaurs and led to the recognition of
two distinctive groups of dinosaurs still current today: Saurischia and Ornithischia (Seeley, 1887). Further
material collected from the Isle of Wight was described by J.W. Hulke and by the late 1880s it became
increasingly obvious that the range and variety of British material attributed to the nominal genus "Iguanodon
mantelli" was disparate stratigraphically and morphologically. Richard Lydekker in a series of review papers
(1888-1890) revised the taxonomy of what was then known and created a new taxonomy that encompassed the
lower and upper Wealden representatives, alongside their known mainland European forms. Recent work that
has not benefitted from the examination of the original material suggests that a radical taxonomic revision is
necessary.

DOLLO, L.,1882. Première note sur les dinosauriens de Bernissart. Bulletin de la Musée Royale d'Histoire Naturelle de Belgique, I: 55-80.
MANTELL, G.A., 1825. Notice on the Iguanodon, a newly discovered fossil reptile, from the sandstone of Tilgate forest, in Sussex.
Philosophical Transactions of the Royal Society of London, CXV: 179-186.
NORMAN, D.B. 1980. On the ornithischian dinosaur Iguanodon bernissartensis from Belgium. Mémoires de l'Institut Royal des Sciences
Naturelles de Belgique, 178: 1-105.
NORMAN, D.B., 1986. On the anatomy of Iguanodon atherfieldensis (Ornithischia: Ornithopoda). Bulletin de l'Institut Royal des Sciences
Naturelles de Belgique, 56: 281-372.
NORMAN, D.B., 1990. A review of Vectisaurus valdensis, with comments on the family Iguanodontidae. In: Carpenter, K. & Currie, P.J.
(eds.), Dinosaur systematics: approaches and perspectives. Cambridge University Press, Cambridge, pp. 147-162.
NORMAN, D.B., 2002. On Asian ornithopods (Dinosauria: Ornithischia). 4. Redescription of Probactrosaurus gobiensis Rozhdestvensky,
1966. Zoological Journal of the Linnean Society (London), 136: 113-144.
NORMAN, D.B., 2004. Basal Iguanodontia. In: Weishampel, D.B, Dodson, P., & Osmolska, H., eds. The Dinosauria. University of
California, Berkeley, pp. 413-437.
SEELEY, H.G., 1887. On the classification of the fossil animals commonly named Dinosauria. Proceedings of the Royal Society of London,
43: 165-171.

Key words: Iguanodon, Wealden, Britain, Europe, taxonomy

74

Darwin-Bernissart meeting, Brussels, February 9-13, 2009

A NEW SAUDI ARABIAN TAXON OF ACANTHOTHORACI
(PLACODERMI): QUESTIONS ON PLATE HOMOLOGIES AND
GROWTH

Sébastien OLIVE 1, 2, Daniel GOUJET 2, Hervé LELIEVRE 2

1 Institut Royal des Sciences Naturelles de Belgique, rue Vautier 29, 1000 Bruxelles,
Sebastien.Olive@sciencesnaturelles.be
2 UMR 5143 CNRS-Muséum National d'Histoire Naturelle de Paris, 8 rue Buffon, 75005 Paris,
goujet@mnhn.fr, lelievre@mnhn.fr

Skull roof patterns of an exceptionally well preserved new Saudi (Al Qalibah area) Acanthothoraci
(Placodermi) are described and submitted to two analysis: the first one aims to establish the skull plates
homologies and the second aims to study their growth process.
This new taxon presents a skull roof subject to an important variability, not only in relation to the
other Acanthothoraci but also in relation to itself; it is illustrated by the presence of supernumerary plates. The
model of placoderm skull roof pattern is generally that of Arthrodira, one of the historically earliest group
described, generally used as a reference for the different taxa of this wide group. The interpretation of the
acanthothoracid skull roof pattern of the Saudi taxon from an arthrodire model is the principal point of this study.
A new plate has been defined: the endolymphatic plate. It is an independent plate, which bears the
endolymphatic foramen. Thanks to that new plate terminology, news homologies can be proposed within the
main groups of placoderms.
Growth concentric lines have been observed on some plates of several specimens. It is the first time
when we can see this kind of lines within the Acanthothoraci. They are particularly well observed on specimens
having suffered a partial erosion. Thanks to this we can analyse the individual growth of the posterior post-
orbital plate, nuchal plate, pre-orbital plate and central plate. All present an allometric growth. During their
development, their relative growth in length is more important than in width. Despite that allometry, we observe
different growth tendencies according to the plate nature and position in the skull roof. Each plate has a
particular behaviour which patterns the overall shape of the skull roof.

Key words: Acanthothoraci, growth, homology, Placodermi, Saudi Arabia

75

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE LATE CRETACEOUS CONTINENTAL VERTEBRATE FAUNA
FROM IHARKÚT, WESTERN HUNGARY A REVIEW OF THE
NEWEST RESULTS

Attila SI

Hungarian Academy of SciencesHungarian Natural History Museum, Research Group for Paleontology,
Ludovika tér 2., Budapest, Hungary, hungaros@freemail.hu

The Late Cretaceous vertebrate locality at Iharkút in the Bakony Mountains (Csehbánya Formation)
yielded the only known Mesozoic continental vertebrate fauna from Hungary. The fluvial Csehbánya Formation
has yielded associated skeletons and over 5000 isolated bones and teeth of at least 24 different vertebrate taxa.
New palynological and paleomagnetic studies further confirmed the earlier established Santonian age of the
bone-bearing beds in the formation. Sedimentological and taphonomical evidences clearly indicate that the
majority of isolated bones, generally collected in fossiliferous pockets, were accumulated in a coarse-grained,
channel deposit of an alluvial flood plain. Along with the remains of invertebrates (ostracods and bivalves),
preliminary results of the stable isotope composition taken from the tooth enamel indicate a freshwater
environment.
Important elements of the fauna are the lepisosteiform and pycnodontiform fishes the latter documented
by jaw elements and teeth, and represent the second known European occurrence of these fishes in freshwater
environment. Besides albanerpetontids, amphibians are represented by two taxa of a new family of frogs. The
toppredators in the aquatic habitat of the Iharkút ecosytem were the mosasaurs (Mosasauridae n. g. n. sp.) which
are known from well-preserved cranial and abundant postcranial material. At least four different taxa of small,
terrestrial lizards are recorded in the assemblage. Remains of turtles indicate the presence of a large-sized
bothremydid turtle whose closest relative is Foxemys mechinorum. Iharkutosuchus makadii was a small-bodied
basal eusuchian crocodylian, up to one metre in length, that is characterized by its heterodont dentition including
multicusped teeth, and a unique transverse jaw movement facilitating effective oral food processing.
Furthermore, the crocodylian assemblage provided evidence for the oldest known alligatoroid eusuchian, and a
ziphosuchian form, Doratodon, which indicates the faunal exchange between Gondwana and the Mediterranean
archipelago.
Pterosaurs were represented by azhdarchids (Bakonydraco galaczi) having a wing span of up to four
metres. The non-avian theropod dinosaur assemblage is composed mainly of isolated teeth and postcranial
remains. Several dozens of medium-sized teeth referred to basal tetanurans are identical with those of the
Campanian of eastern Austria. Although another non-avian theropod taxon is known only from a single, well-
preserved ungual phalanx, its characters unambiguously refer it to the Abelisauridae. This find is the oldest
occurrence of the group in Europe and it further strengthens the paleogeographic argument that besides
Laurasian taxa, the Mediterranean archipelago was also inhabited by Gondwanan forms already in the early Late
Cretaceous. The richest material of non-avian theropods is from small-bodied dromaeosaurs. Iharkút is among
the exceptional localities where avian material has been also found. The few isolated limb bones indicate at least
two different taxa of Enantiornithes. In contrast to most other Late Cretaceous vertebrate sites in Europe,
rhabdodontid ornithopods are poorly known in Iharkút. The best represented archosaurian taxa is the nodosaurid
ankylosaur, Hungarosaurus tormai. Based on comparative anatomical and phylogenetic analyses, several taxa,
reported from the Santonian of Iharkút show a close relationship with much older forms. The closest relative of
Iharkutosuchus is Hylaeochampsa vectiana from the Barremian of the Isle of Wight. The basal tetanuran teeth
from Iharkút strongly resemble the teeth of M. dunkeri from the Wealden of England, and Hungarosaurus
(together with Struthiosaurus) are more primitive than their Early to Middle Cretaceous relatives. These features
of the fauna may suggest that the Iharkút area existed as a refugium during the Santonian.

Key words: vertebrate fauna, Santonian, Iharkút, Hungary

76

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
A NEW IGUANODONTIAN DINOSAUR FROM THE EARLY
CRETACEOUS YIXIAN FORMATION OF WESTERN LIAONING,
CHINA

Rui PAN 1, Yihan WANG 2, Xiaolin WANG 1

1 Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044,
China, P.O.Box 643 deep.bluesea@163.com
2 Department of Biology, Capital Normal University, No.105 Northwest Third Ring Road, Beijing 100048,
China

In this paper we will report a new juvenile iguanodontian dinosaur from Yixian Formation, west
Liaoning. China. So far a lot of feathered theropod and small ceratopsian dinosaurs have been reported, only
quite a few large herbivorous dinosaurs such as Jinzhousaurus are known in Yixian Formation. Here we describe
a new species of large herbivorous dinosaur. Some characters are different from that of the reported ones. The
premaxilae, unlike that of Jinzhousaurus or Iguanodon bernissartensis, the rostral portion doesn't prominent
curve downward to the tip of the snout. And they do not expande laterally to produce a flared "beak". The
surangular, like that of Altirhinus and Lanzhousaurus, there is a small embayment present at the lever with the
base of the coronoid process, but such embayment is absent in Equijubus and Iguanodon.

The humeri are straight and have low elto-pectoral crest. That aparently differ with that of
Jinzhousaurus, Hadrosauridae or Nanyangosaurus, but similar with Camptosaurus and Ouranosaurus.

The illum has a long tape preacetabular process with a slightly everted dorsal margin. The preacetabular
process forms a "u" shape with the public peduncle. The pubic peduncle curves smoothly to the ischium
peduncle. These are much like Ouranosaurus nigeriensis but quite different with Iguanodon and Camptosaurus.

The shaft of the pubis is nearly half of the length of ischium, that's quite different with Dryosaurus
altus and Tenontosaurus tilletti, while their pubis has almost the same length of ischium. And its prepubic
process is blade-shaped and expands distally, taping slightly toward its distal end. But the anterior part of the
blade doesn't upturn. It also lacks a symphysis. These are similar characters with Ouranosaurus and
Hadrosaurids.

Key words: Iguanodontia, Jehol Biota, Early Cretaceous, Yixian Formation, west Liaoning

77

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
LOWER CRETACEOUS DINOSAURS OF SPAIN: AN OVERVIEW
BASED ON SKELETAL REMAINS

Xabier PEREDA SUBERBIOLA 1, José Ignacio RUIZ-OMEÑACA 2,3,
José Ignacio CANUDO 3, Fidel TORCIDA 4, José Luis SANZ 5

1 Universidad del País Vasco/EHU, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y
Paleontología, Apartado 644, 48080 Bilbao, Spain, xabier.pereda@ehu.es
2 Museo del Jurásico de Asturias (MUJA), 33328 Colunga, Spain, jigruiz@unizar.es
3 Grupo Aragosaurus (www.aragosaurus.com), Paleontología, Facultad de Ciencias, Universidad de
Zaragoza, Pedro Cerbuna 12, 50009 Zaragoza, Spain, jicanudo@unizar.es
4 Colectivo Arqueológico-Paleontológico de Salas (C.A.S.), Museo de Dinosaurios, Plaza Jesús Aparicio 9,
09600 Salas de los Infantes, Burgos, Spain, secretaria@colectivosalas.com
5 Unidad de Paleontología, Departamento de Biología, Universidad Autónoma de Madrid, c/Darwin 2, 28049
Cantoblanco, Madrid, Spain, dinopepelu@gmail.com

The Lower Cretaceous formations of Spain (mainly the Wealden facies [Valanginian-Barremian] and
those correlated with the Urgonian deposits [Barremian-Aptian]; the Purbeck facies [Tithonian-Berriasian],
which include the Jurassic-Cretaceous transition, are not considered here) have yielded abundant dinosaur
remains. The fauna consists of theropods, sauropods, thyreophorans and ornithopods; seven genera and species
defined in Spain are exclusively known in this country. The Spanish record is one of the best known in Europe,
only surpassed in biodiversity by the classical dinosaur sites of England. Although the earliest dinosaur remains
from the Early Cretaceous of Spain were described in the 1870s, the first fieldworks were not undertaken until
the 1920s. Most of the meaning discoveries have been made in the last 25 years.
The most significant Spanish locality is the Konservat-Lagerstat of Las Hoyas (Cuenca). Othe important
sites include those of Salas de los Infantes (Burgos), Galve (Teruel), Morella and Cinctorres (Castellón).
Golmayo (Soria), Igea (La Rioja), El Montsec (Lleida), Castellote, Josa and Peñarroya de Tastavins (Teruel),
and Uña (Cuenca) have also yielded relevant dinosaur specimens. With the exception of El Montsec (Pyrenean
Realm), all the others mentioned belong to the Iberian Chain (Cameros Basin, Maestrazgo Basin, Central and
SW Iberian Chain). Dinosaur tracksites, very abundant in the Cameros Basin, are not taken into account here.
Ornithopods are the most abundant and diversified dinosaurs from the Early Cretaceous of Spain.
Iguanodontoids consists of Iguanodon bernissartensis and Iguanodon (Mantellisaurus) cf. atherfieldensis from
Castellón, Cuenca, Teruel and Burgos, and a new genus and species yet undescribed from Teruel. Other Spanish
material is provisionally referred to as Iguanodon sp. or as Iguanodontoidea indet. "Hypsilophodontids" are
represented by Hypsilophodon-like and Othnielia-like forms from Burgos, Castellón, La Rioja and Teruel.
Dryosaurids includes Valdosaurus sp. and "Camptosaurus" valdensis from Burgos and Teruel.
Among theropods, the ornithomimosaur Pelecanimimus polyodon and the enantiornithine birds
Iberomesornis romerali, Concornis lacustris, Eoalulavis hoyasi are known from Cuenca, and the enantiornithine
Noguerornis gonzalezi from Lleida. Indeterminate enantiornithine specimens have also been found in the same
sites. Spinosauroids include Baryonyx-like and indeterminate baryonychine material from Burgos, La Rioja,
Teruel and Castellón. In addition, other theropod taxa have been described in the Spanish record: allosauroids,
carcharodontosaurids, dromaeosaurids, Paronychodon-like and Richardoestesia-like, most of them on the basis
of isolated teeth.
The sauropod association is dominated by titanosauriforms. Tastavinsaurus sanzi and tentatively
Aragosaurus ischiaticus are regarded as basal somphospondylians. The presence of a member of Titanosauria in
Teruel needs to be confirmed on the basis of more complete material. Basal titanosauriforms are represented by
indeterminate brachiosaurids from Castellón and "Pleurocoelus" valdensis teeth from Teruel. Teeth similar to
those of Euhelopus have also been described in Teruel.. The only non-titanosauriform sauropods are a
rebbachisaurid diplodocoid from Burgos (that probably represents a new genus and species) and teeth of
"Oplosaurus armatus" from Teruel.
Finally, the thyreophorans consists of the ankylosaur Polacanthus sp. from Burgos, Castellón and Soria,
and indeterminate stegosaurs from Burgos and Teruel.
The dinosaur associations of the Early Cretaceous of Spain are singular and complex. These faunas are of
particular palaeobiogeographical interest on account of the special location of the Iberian Plate to provide a
better knowledge of the connections between the Laurasian and Gondwanan landmasses during the Early
Cretaceous.

Key words: Dinosaurs, Early Cretaceous, Valanginian-Aptian, Spain

78

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EARLY CRETACEOUS ORNIHTOPOD DINOSAURS FROM ROMANIA

Erika POSMOSANU

Tarii Crisurilor Museum, B-dul Dacia nr. 1-3, 410464 Oradea, Romania,
eposmosanu@gmail.com

The Early Cretaceous (Berriassian-Valanginian) vertebrate fauna from the bauxite deposits of Cornet
Lens 204, North-Western Romania consists of dinosaurs, pterosaurs and possibly birds. Among dinosaurs,
ornithopods are over-represented. The fragmentary, disarticulated, and somewhat sorted nature of the material
makes establishing the taxonomic identities and ontogenetic age difficult. Tiberiu Jurcsak made the first
taxonomic interpretations of the ornithopod fauna and identified Hypsilophodon sp., Valdosarurus
cannaliculatus, Iguanodon cf. mantelli, and Vectisaurus valdensis (Jurcsak and Popa, 1978, 1979). The latter
was subsequently considered a junior synonym of Iguanodon atherfieldensis (Norman, 1990). Revision of the
ornithopod fauna concluded that Hypsilophodon can not be attested, the Valdosaurus remains can be identified
as Dryosauridae indet., among Iguanodontidae the material is referable to Iguanodon sp., and among
euornithopoda, to Camptosaurus sp. (Tallodi-Posmosanu and Popa, 1997; Posmosanu, 2003).
Ornithopod postcranial anatomy is fairly conservative, so most of the Cornet material cannot be
taxonomically identified with confidence. Norman and Barrett, 2002 compared Camptosaurus and Iguanodon
teeth and referred I, hoggii to as Camptosaurusi. Tallodi-Posmosanu and Popa (1997) first recognised the
similarities between the Cornet dentary teeth and those of C. hogii. The lingual surface of the Cornet dentary
teeth crowns (MTCO [Tarii Crisurilor Museum Oradea] 14.333, 19.195, 20.265, 20.796, 20.798) are equally
sub-divided by simple, well-defined and equal sized primary and secondary ridges, tertiary ridges are absent.
The lingual surface of each crown is covered by a thick layer of enamel, which exhibits visible transverse
fluting. The consistent feature seen in the dentitions, which separates Camptosaurus hoggii from any other taxa
according to Norman and Barrett (2002) is the absence of an abbreviated cingulum on the distal margin of the
crown. None of the Cornet dentary teeth crowns displays this distinctive inrolled edge on the distal margin of the
crowns, resembling the abbreviated cingulum. The ornamentation of the marginal denticles in the best preserved,
unworn teeth resembles the ornamentation described by Norman and Barrett (2002) for C. hogii. Thus, the
Cornet dentary teeth can be referred to Camptosaurus hoggii. The systematic position of C. hoggii was
considered at best provisional because the holotype does not possess unambiguous diagnostic features (Norman
and Barrett, 2002). According to Carpenter and Wilson (2008), C. hoggii does not belong to the genus
Camptosaurus, because its dentary is too different to that of C. aphanoecetes and C. dispar.
Because of the isolated, fragmentary nature of the fossils and the fact that some postcranial elements of
Dryosauridae are not distinguishable from those of small camptosaurs and small iguanodontids, the Cornet
material can be most confidently identified as a medium-sized, camptosaur-like ornithopod, living near Cornet in
Berriassian Valanginian period.

CARPENTER, K, & WILSON, Y., 2008. A new species of Camptosaurus (Ornithopoda: Dinosauria) from the Morrison Formation (Upper
Jurassic) of Dinosaur National Monument, Utah and a biomechanical analysis of its forelimb. Annals of Carnegie Museum, 76: 227-
263.
JURCSÁK, T., POPA, E., 1978. Resturi de dinosaurieni în bauxitele de la Cornet (Bihor). Nymphaea, 6: 61-64.
JURCSÁK, T., POPA, E., 1979. Dinosaurieni ornitopozi din bauxitele de la Cornet (Munii Pdurea Craiului). Nymphaea, 7: 37-75.
NORMAN, D.B., 1990. A review of Vectisaurus valdensis, with comment son the family Iguanodontidae. In: Carpenter, K. & Currie, P.J.
(eds.), Dinosaur Systematics: Approaches and Perspectives. Cambridge University Press, Cambridge, pp.147-161.
NORMAN, D.B. & BARRETT, P.M., 2002. Ornithischian dinosaurs from the Lower Cretaceous (Berriassian) of England. Special Papers in
Palaeontology, 68: 161-189.
POSMO ANU, E., 2003. Revision of the Early Cretaceous dinosaur (ornithopoda) collection from the bauxite deposit Lens 204 - Cornet,
Romania. Nymphaea, Folia Naturae Bihariae, 30: 25-38.
TALLODI-POSMOSANU, E. & POPA, E., 1997. Notes on a Camptosaurid dinosaur from the Lower Cretaceous bauxite, Cornet
Romania. Nymphaea, 23-25: 35-44.

Key words: Early Cretaceous, Ornithopoda, Camptosaurus, Romania

79

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
FAUNAL ASSEMBLAGES AT THE JURASSIC/CRETACEOUS
BOUNDARY: COMPARISON BETWEEN SOME WEST EUROPEAN
CONTINENTAL SITES

Joane POUECH, Jean-Michel MAZIN

Université Claude Bernard Lyon 1, UMR 5125 PEPS,
La Doua Bât. Géode 2, rue Dubois, 69622 Villeurbanne, France
joane.pouech@pepsmail.univ-lyon1.fr
jean-michel.mazin@univ-lyon1.fr

Study of the Berriasian site of Cherves-de-Cognac (Charente, France) has revealed a rich and
diversified vertebrate fauna, mainly known from dental micro-remains. This fauna was deposited in a lagoon
environment, evolving from brackish water to lacustrine deposits. Quantitative study of the biodiversity from
micro-vertebrates leads to understand the depositional modalities of this allochtonous fauna brought in the
depositional setting. Three families of aquatic vertebrates are well represented, associated with amphibious taxa
(crocodilians). The terrestrial fauna is recorded only occasionally, thanks to exceptional events as flood or
continental washing. Among this fauna, mammals are well represented as they contribute for more than 35% of
the terrestrial diversity.
The vertebrate fauna identified is close to other West European records from diverse continental or
margino-littoral environnements, and it is interesting to replace the information from Cherves-de-Cognac in its
temporal and geographical context. Comparison includes West European sites ranging from the Kimmeridgian to
the Barremian, which have yielded a well diversified fauna and/or Mesozoic mammals, known for macro- and/or
micro-remains: Bornholm (Denmark), Bernissart (Belgium), Isle of Wight and Purbeck Limestone Group
(England), Boulonnais, Canjuers, Cerin and Chassiron (France), Oker and Solnhofen (Germany), Buenache,
Galve, Las Hoyas, Montsec, Uña and Vallipón (Spain), Guimarota and Lourinhã Group (Portugal).
Multivariate analyses regroup the sites according to the ecology (aquatic or terrestrial) of the main fauna
recorded. Differences in depositional environment or age are of weak influence in the organisation of the
phenograms. On another hand, it seems there is no discrimination based on the geographic distribution of the
sites.
Moreover, this analysis reveals a hierarchy in the recording of the continental fauna in these
assemblages. Aquatic and amphibious taxa are first deposited in the depositional setting. Then, theropods are
currently known, sometimes with others families of dinosaurs. It seems that mammals are present only if all the
precedent members are already known in the fossil assemblage.
Finally, the distribution of the mammals families around the Jurassic/Cretaceous boundary suppose a
faunal change among the mammalian community, which occurs probably during the Berriasian, which questions
the positioning of the limit between the two systems.

Key words: Jurassic/Cretaceous boundary, West European continental sites, faunal assemblages

80

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
DESCRIPTION AND WEST EUROPEAN AFFINITIES OF THE
MAMMALIAN FAUNA OF CHERVES-DE-COGNAC

Joane POUECH, Jean-Michel MAZIN

Université Claude Bernard Lyon 1, UMR 5125 PEPS,
La Doua Bât. Géode 2, rue Dubois, 69622 Villeurbanne, France
joane.pouech@pepsmail.univ-lyon1.fr
jean-michel.mazin@univ-lyon1.fr

The site of Cherves-de-Cognac (Berriasien, Early Cretaceous, Charente, France) has yielded a
diversified and abundant fauna, mainly known from dental micro-remains. Several teeth of mammals have been
found corresponding to six families, present in West Europe at the Jurassic/Cretaceous boundary.
Three teeth belong to the order Eutriconodonta, two of them are reported to the sub-family
Triconodontinae. Multituberculates are known by a unique species, Pinheirodon pygmaeus, represented by
several dental morphologies (upper incisors, upper and lower premolars, lower molar). Two families of
symmetrodonts have been identified. The first one, Thereuodontidae, is known by an upper molar, but the
validity of this taxon can be questioned. Three teeth are reported to the family Spalacotheriidae, among them one
molar is close to the species Spalacotherium evansae. The family Dryolestidae is the one of the best represented
family, known by an upper molar and several lower molars. Finally, a half lower molar (trigonid) is conferred to
the genus Peramus.
Terrestrial fauna is known in an unique level in the sedimentary series of Cherves-de-Cognac, where
mammals represent only 0.08% of the vertebrate specimens, but 19% of the family diversity. Among these
terrestrial taxa, mammals contribute for more than 35% of the number of specimens and number of families. The
Cherves-de-Cognac mammalian assemblage is mainly represented by multituberculates Pinheirodontidae (50%
of the known specimens) and Dryolestidae (23%).
By comparison with the others west european sites, the identified assemblage suggest closer affinities
with the fauna from the Upper Jurassic and the Early Cretaceous, especially with Porto Pinheiro (Berriasian,
Portugal) and particularly with the Purbeck Limestone Group (Berriasian, England), where the six families from
Cherves-de-Cognac are already known.

Key words: Cherves-de-Cognac, Berriasian, Mesozoic mammals

81

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE KARSTIC PHENOMENON OF BERNISSART PIT AND THE
GEOMORPHOLOGIC SITUATION IN THE MESOZOIC TIMES

Yves QUINIF

Faculté Polytechnique de Mons, rue de Houdain, 9, 7000 Mons, yves.quinif@fpms.ac.be

Bernissart pit belongs to a family of great vertical cylindrical features which cross over the
Carboniferous terranes of Pennsylvanian and have their roots in the carbonated Mississippian. Their heights are
more of some hundred meters. Their origin is due to karstic phenomena which developed in the Mississippian
carbonates and evaporites. Some pits are blind: their summits are situated in the coal bearing formations.
Geological observations have permitted to find any pits which have reached the summit of the Paleozoic
basement. In this case, the upper part of the pit is filled by Cretaceous sediments which are dated from
Barremian to Albian (in Bernissart pit) or Turonian (in Ghlin pit).

The genesis of those pits depends of the intensity and the dating of the deep karstification of the
carbonated and evaporitic Carboniferous. Thus, the ages of the trapped sediments give an upper limit for that
deep karstification. We conclude that this karstification has had its intensity maximum from the lower to the
beginning of the upper Cretaceous. During this times interval, the limestone's at the north border of the Mons
Basin from Tournai to Soignies are well karstified. Thus, we note that an intense phase of karstification takes
place during this period. We can also conclude that the voids which contain presently hot water in the
geothermal aquifer develop at this time.

Two important problems must be solved: why this karstification phase takes place during this period
and how can we reconcile dissolution at a precise point which can generate a high karstic pit and a regional
dissolution which can conduct to a regional subsidence. To have a deep karstification, we must have an input and
an output of the aggressive fluids. The input region can be localized at the north of the Mons Basin where
limestones outcrop. But the output area remains obscure. The deep karstification is essentially provoked in this
time interval because there is a tectonic phase in extension, which permits subsurface karstification at the North
and the deep penetration of aggressive water under the Pennsylvanian. The output of the saturated solutions
would can possible by convection cells in the thickness of the Mississippian. The stop of the karstification
situated when the great transgressions set up, during the Turonian. Concerning the relation between dissolution
at a precise point which can generate a high karstic pit and a regional dissolution, the origin could be in a
beginning dissolution process at a precise point, controlled by tectonics conditions. After, following some
favorable bed, this dissolution could progress laterally.

CORNET J. & BRIARD A., 1870 Notice sur les puits naturels du terrain houiller. Bull.Soc.Belg.Géol., Pal. Et d'Hydr., XXIX, n°5.
DELMER A., LECLERQ V., MARLIÈRE R., ROBASZYNSKY F., 1982 La géothermie en Hainaut et le sondage de Ghlin (Mons,
Belgique). Ann.Soc.Géol. du Nord, TCI : 189-206.
DELMER A. & VAN WICHELEN P., 1980 Répertoire des puits naturels connus en terrains houiller du Hainaut. Prof. Paper, Service
Géologique de Belgique, 172 p.
DUPUIS C. & VANDYCKE S., 1989 Tectonique et karstification profonde : un modèle de subsidence original pour le Bassin de Mons.
Ann.Soc.Géol.Belg., 112, 2 : 479-488.
MARLIÈRE R., 1932 Une roche peu connue provenant d'un puits naturel de la région de Bernissart. Ann.Soc.Géol.Belg., 55 : 184-189.
QUINIF Y., 1995 Le Puits de Flénu (Belgique). La plus grande structure endokarstique au monde (1200 m) et la problématique des puits
du Houiller. Karstologia, 24 : 29-36.
QUINIF Y., VANDYCKE S., VERGARI A., 1997 - Chronologie et causalité entre tectonique et karstification - l'exemple des paléokarsts
crétacés du Hainaut (Belgique). Bull.Soc.Géol.Fr., 168, 4 : 463-472.
VERGARI A., QUINIF Y., 1997 Les paléokarsts du Hainaut. Geodinamica Acta. 10, 4 : 175-187.

Key words: Bernissart, pits, karstification

82

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
PRESENCE OF SEBECOSUCHIAN CROCODYLIFORMS IN THE
LATE CRETACEOUS OF EUROPE

Márton RABI

Department of Paleontology, Eötvös Lorand University, 1117 Pázmány Péter st. 1/C. Budapest, Hungary,
iszkenderun@freemail.hu

Sebecosuchian crocodiles are bizarre, deep snouted, ziphodont forms well adapted for terrestrial,
carnivorous lifestyle. Most of their record comes from the Upper Cretaceous Miocene of South America and
only a few remains have been reported from the Late Cretaceous of Pakistan and the Eocene of Portugal, Spain
and France (Ortega et al., 1996; Carvalho et al., 2005; Paolillo & Linares, 2007). Questionable sebecosuchains
are known from the Eocene of Algeria and Germany, but their affinities with the Trematochampsidae can not be
excluded (Buffetaut, 1988). The Sebecosuchia is widely accepted to form a monophyletic group and together
with its presumed sister-taxon, the Notosuchia, they make up the larger clade Ziphosuchia.
The Late Cretaceous crocodile fauna of Europe is dominated by eusuchains, no sebecosuchians have been
identified before. The only described non-eusuchian crocodiles are Ischyrocampsa from the Campanian of
France and Doratodon from the Campanian of Austria and Spain. Ischyrocampsa is considered to be a
trematochampsid and Doratodon is here recognized as a sebecosuchian. Presently, Doratodon comprises two
species, the type species Doratodon carcharidens Bunzel, 1871 from Austria and Doratodon ibericus Company,
Suberbiola, Ruiz-Omenaca & Buscalioni, 2005 from Spain. As Doratodon is mainly known on the basis of
mandibular material (only part of the maxilla has been known from the skull) its phylogentic relationship has
been highly controversial during the last two centuries. The most recent study was that of Company et al. (2005)
who performed a phylogenetical analysis which placed Doratodon as the sister-taxon of the Sebecosuchia. The
study of new Doratodon material from the Santonian of Hungary (Iharkút locality) and the revision of the type
of D. carcharidens revealed that this taxon is in fact lies within the Sebecosuchia. Comparisons to several non-
neosuchians, including the Notosuchia, Trematochampsidae, Peirosauridae and Araripesuchidae were made
mainly using the literature and the most similarities were found with sebecosuchians. The relationships of
Doratodon within this group were investigated for the first time.
The Hungarian occurrence of the genus is represented by a fragmentary maxilla and a dentary and isolated
teeth. A pterygoid and a quadrate are also referred to this form. Comparisons between the two species of
Doratodon showed that the Hungarian taxon is more closely related to D. carcharidens than to D. ibericus.
Concerning the pterygoid from Iharkút Doratodon is more primitive than the Baurusuchidae and Bretesuchus
and propably more closely related to Sebecus, Zulmasuchus and Barinasuchus because of the participation of the
pterygoid in the suborbital fenestra and the absence of an extra pterygoid foramen. Doratodon seems to be more
primitive than these latter three because of the presence of an antorbital fenestra but it is probably more derived
than Pehuenchesuchus because of the presence of serrated teeth.
The new interpretation of Doratodon implies that the dispersion of the group to Europe was not through
Indopakistan and Asia during the Eocene as previously suggested by Carvalho et al. (2005). It is most likely that
Sebecosuchians used the South-American-African landbridge which persisted at least until the Aptian and they
reached Europe from North-Africa through Apulia and Iberia at least by the Santonian. This hypothesis is
consistent with new discoveries from Africa and also with the presumed mid-Cretaceous apperaence of the
Sebecosuchia as suggested by numerous ghost lineages. However, it supposes a later derivation of the group than
the vicariant model of Turner (2004).

BUFFETAUT, E., 1988. The Ziphodont Mesosuchian Crocodile from Messel: a reassessment. Cour. Forsch.-Inst. Senckenberg, 107: 211-
221.
CARVALHO, I. S., CAMPOS., C. A., & NOBRE, P. H., 2005, Baurusuchus salgadoensis, a new Crocodylomorpha from the Bauru Basin
(Cretaceous), Brazil. Gondwana Research, 8: 11-30.
COMPANY, J., SUBERBIOLA, X. P., RUIZ-OMENACA, J. I. & BUSCALIONI, A. D., 2005: A new species of Doratodon
(Crocodyliformes: Ziphosuchia) from the Late Cretaceous of Spain. Journal of Vertebrate Paleontology, 25: 343-353.
ORTEGA, F., BUSCALIONI, A.D., & GASPARINI, Z., 1996. Reinterpretation and new denomination of Atacisaurus crassiproratus
(Middle Eocene; Issel, France) as cf. Iberosuchus (Crocodylomorpha, Metasuchia). Geobios, 29: 353-364.
PAOLILLO, A. & LINARES, O. J., 2007. Nuevos crocodilos Sebecosuchia del Cenozoico Suramericano (Mesosuchia: Crocodylia).
Paleobiologia Neotropical, 3: 1-25.
TURNER, A. H., 2004. Crocodyliform biogeography during the Cretaceous: evidence of Gondwanan vicariance from biogeographical
analysis. Proc. R. Soc. B, 271: 20032009.

Key words: Crocodiles, Sebecosuchia, Europe, Hungary, Late cretaceous

83

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
FORAMINIFERA FROM THE BERNISSART BORING F3
IN THE CONTEXT OF THE MID-CRETACEOUS TRANSGRESSION

Francis ROBASZYNSKI

Faculté Polytechnique de Mons, 9 rue de Houdain, 7000 MONS, francis.robaszynski@fpms.ac.be

The Bernissart well F3 was bored in 2002 through the whole Cretaceous-Paleogene succession of the
Mons basin to locate the levels of the wealden facies containing the famous Iguanodons of Bernissart. The
description of the drilling cores gave the following lithological units (Yans, 2007):

0-8m: glauconitic sands (Paleogene, Thanetian),
8-75m: glauconitic white chalk, chalk with flints and cherts, marls, basal glauconitic conglomerate = Tourtia de
Mons (Middle Cretaceous),
75-265m: sandstones, sandy calcarenites, numerous sponge spicules, conglomerates (Albian "meule"),
265-315m: black clays and clayey sands (wealden facies, Barremian),
315-349m: breccias, fragments of Namurian rocks in a clayey matrix.

The goal of the present study was to have more details on the Cretaceous stratigraphic succession. From
7 to 245.50m, 52 samples were taken and processed to release the foraminiferal content, from which 38
delivered benthic and/or planktonic forms. The main biostratigraphic results are summarized in what follows:

245.50 to 189.40m: Upper Albian sensu stricto by the association of Arenobulimina chapmani Cushm.,
Orithostella jarzevae (Vass.), A. cf. sabulosa (Chapm.), Gavelinella cenomanica (Brotz).
109.60 to 103.10m: Vraconnian by the association of G. baltica (Brotz.), Epistomina cf. cretosa/spinulifera
(Reuss), Vaginulina strigillata bettenstaedti Albers,
77.50 to 70.60m: Cenomanian/Turonian transition with the association of G. cenomanica, G.baltica,
Lingulogavelinella globosa (Brotz.),
69.50 to 26.20m: Turonian by the association of Whitenella archaeocretacea Pess., Praeglobotruncana stephani
(Gand.), Dicarinella hagni (Scheibn.), Bdelloidina cribrosa (Reuss), Helvetoglobotruncana helvetica (Bolli),
Marginotruncana sigali (Reich.), M. pseudolinneina Pess., Globorotalites micheliniana (d'Orb.) and the
mesofossil Terebratulina rigida,
24.80 to 8m: Coniacian by the association Gavelinella arnagerensis Solakius, Reussella kelleri Vass.,
Stensioeina granulata granulata (Olbertz), Globotruncana linneiana (d'Orb.).
A distribution chart of all species of foraminifera found in the F3 boring will be presented in the poster.

It is to note the lack of the uppermost Cenomanian as any Rotalipora cushmani was found below the
basal Turonian conglomerate ("Tourtia de Mons"). The same feature was observed at the boring Bernissart 41
(Robaszynski, 1972a). The two borings Bernissart F3 and 41 where the Upper Cenomanian marls lack are
situated at the marginal northern part of the Mons basin when, in the central part, at Trith near Valenciennes as
in the southern part near Bettrechies and Maubeuge, the Cenomanian transgression leaved several metres of
marls containing R. cushmani (Robaszynski, 1972b; Leplat & Robaszynski, 1972).

ROBASZYNSKI F. (1972a).- Les foraminifères pélagiques des « Dièves » aux abords du golfe de Mons (Belgique). Annales de la Société
géologique du Nord, 91, 31-38.
ROBASZYNSKI F. (1972b).- Les « Dièves » de Maubeuge et leurs deux Tourtias (Crétacé supérieur). Annales de la Société géologique du
Nord, 91, 193-197.
LEPLAT J. & ROBASZYNSKI F. (1972).- Une couche à Rotalipores dans les "Dièves" (Crétacé supérieur) dans un sondage à Trith (Nord).
Annales de la Société géologique du Nord, 91, 199-202.
YANS J. (2007).- Lithostratigraphie, minéralogie et diagenèse des sédiments à faciès wealdien du Bassin de Mons (Belgique). Académie
royale de Belgique, mémoire in 4° de la Classe des Sciences, t.9, n° 2046, 178p.
YANS J., DEJAX J., PONS D., TAVERNE L. & BULTYNCK P. (2006).- The Iguanodons of Bernissart are middle Barremian to earliest
Aptian in age. Bulletin de l'Institut royal des Sciences naturelles de Belgique, 76, 91-95.

Key words: Bernissart, Foraminifera, `mid' Cretaceous

84

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE ATTEMPTED THEFT OF DINOSAUR SKELETONS DURING THE
GERMAN OCCUPATION OF BELGIUM 1914-1918 AS A TOPIC OF
THE HISTORY OF SCIENCE AND THE HISTORY OF TOTAL WAR

Christoph ROOLF, M.A.

Heinrich-Heine-Universität Düsseldorf, Historisches Seminar II (Neuere Geschichte), Universitätsstraße 1,
D 40225 Düsseldorf, Germany, roolf@uni-duesseldorf.de

My presentation deals with the hitherto largely unknown attempts of German scientists to seize Belgian
cultural possessions during the First World War. Probably the most spectacular examples of these are the
activities of German palaeontologists and German natural history museums at the biggest dinosaur excarvation
site in Europe, which is located in the Belgian town of Bernissart. These took place during the German
occupation of the country between 1914 and 1918. Following a plan of the German paleontologist Otto Jaekel
(launched in spring 1915) another Iguanodon skeletons were to be excarvated and transferred to German natural
history museums. The work began in June 1916 and ended without results with the retreat of German troops
from Belgium in late 1918.

The establishment of German occupation authorities in large parts of Western and Eastern Europe in the
late summer of 1914 opened up new and unexpected fields of activity for scientists and scientific institutions.
This included direct participation in the governing bodies as well as counselling functions which were to support
German occupation policies and war aims. Most of all, the occupied territories offered ample opportunities to
conduct ambitious research projects with the cooperation of the occupation authorities. This would often lead to
a loot of cultural possessions. That the German occupation was actually a necessary precondition for some
research projects is illustrated by the following considerations:

To analyse scientific activities during war time it is necessary and helpful to combine different
approaches. Analytical tools from the history of science can be used in connection with those from the history of
particular fields of science and general history, with special emphasis on the analysis of modern societies in
times of total war. This is of crucial importance because it is the more or less complete exploitation of the
civilian and material resources of occupied countries that is the hallmark of total warfare in the 20th century.

The case of the excarvation site in Bernissart during the First World War shows that even a seemingly
unpolitical, scientific project can become a battleground for different (and often opposing) interest groups within
the occupying body. The driving force behind the plan to further excarvate and ,,export" dinosaur skeletons from
Bernissart was the paleontologist Otto Jaekel who was a professor at Greifswald university and had come to
Brügge in Belgium in the spring of 1915 as part of a reserve regiment. By proposing new excarvations he was
primarily hoping to further his own career. The second major benefactor from the project was going to be the
Museum of Natural History in Berlin. It was to receive most of the expected fossils and thus to gain prestige
among the leading museums of natural history in the world. The scientific museums and collections in Munich,
Frankfurt, Hildesheim and Hamburg also hoped for a boost in their reputation by contributing to costs of the new
excarvations. In return, they were to receive some Iguanodons of their own. The occupation administration in
Brussels, German authorities and Prussian ministeries in Berlin as well as the German emperor, they all endorsed
the plans for new excarvations as long as it did not affect other war aims in Belgium. Thus, the occupation
authorities strictly refrained from breaking the local resistance against the new excarvations by force. This
resistance originated mainly from the coal mines in Bernissart, where the digging was scheduled to take place. It
was also resisted by the Museum of Natural History in Brussels. Jaekel, however, had urged for a German
administrative body to be superimposed on the mines to enforce his plan. This idea that was strongly rejected by
the occupying authorities who clearly saw the importance of Belgian coal for the German war effort and who
were also worried about unrest among the workers and international protests against any action that could be
construed as another case of ,,cultural barbarism" of the Prussian militarism.

Key words: Belgium, dinosaurs, First World War, German occupation policy, plundering cultural assets

85

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE RECORD OF CLIMATE AND EUSTATIC CHANGES DURING
THE LOWER CRETACEOUS IN THE "ARCILLAS DE MORELLA
FORMATION" (SPAIN)

Carlos de SANTISTEBAN 1, Andrés SANTOS-CUBEDO 2,3

1 Departament de Geologia, Universitat de València, E-46100- BURJASSOT (València), Spain,
Carlos.santisteban@uv.es
2 Departament de Recerca del Mesozoic, Institut Català de Paleontologia (ICP), c/Escola Industrial, 23, E-
08201 Sabadell (Barcelona), Spain, andres.santos@icp.cat
3 Grup Guix, c/ Santa Lucia 75, E-12540 Vila-real (Castellón), Spain

The "Arcillas de Morella" formation is a clastic unit that contains dinosaur remains - among them those of
Iguanodon benissartensis (Santafé, et al. 1982) -, from the Maestrat Mesozoic basin. This basin was part of the
northern margin of the Tethys and was placed in the eastern sector of the Iberian plate during the Lower
Cretaceous.
The "Arcillas de Morella" formation has a maximum thickness of 90 m and is early middle Aptian in
age. It is formed by red clays, sandstones, conglomerates, green marls and limestones. In the geologic literature,
the materials of this unit have been interpreted as continental deposits formed in fluvial floodplain environments
of a tidal-dominated delta. However, a detailed analysis of the fauna found in the localities with dinosaur fossils
and the geometry of the sedimentary bodies allow thinking about a new interpretation.
Most of the deposits of this unit are constituted by red clays with intercalations of green marls forming
cycles of marine (marls) and continental (red clays) materials. The sandstones are present in big channelled
bodies or forming laterally extensive units. The second ones adapted to erosion surfaces. The channels are 700 m
in cross section and up to 25 m of depth. In them, besides the sandstones, there are conglomerates, green marls
and carbonates that contain marine fauna (foraminifers, oyster shells and echinoids) and remains of trunks with
borings of marine bivalves. Locally the channels include very thin laminate facies, with alternations of silt and
dark grey lime, rich in organic mater without bioturbation traces. Most of the localities with dinosaur fossils in
this formation are in the infilling parts of the channels. In these sites remains in anatomical connection have been
discovered, but also reworked bones that, in some cases, they contain autochthonous marine fauna.
Many of the sandstones bodies are beach deposits and they are located inside incised valleys. They
constitute the transgressive system tract of the cycles represented by the alternations of green marls and red
clays. These cycles contain carbonates with tropical marine fauna, at the base, and they finish, on top, in a
calcareous paleosoil similar to the modern caliches developed in semi-arid Mediterranean regions. Hence these
eustatic cycles are in connection with climatic changes developed in the western Tethys during the early
Cretaceous. Cycles of the same type have also been described in the sedimentary record of the upper Jurassic of
the eastern Iberia (Santisteban, 2004) and they can be recognized with independence of the tectonic control.
The existence of climatic changes accompanied by eustatic ones in the early Cretaceous in the northern
margin of Tethys, in the eastern Iberia, should have affected the composition and structure of the terrestrial
ecosystems. The presence of marine and continental fauna, and autochthonous and allochthonous remains in the
same fossil sites, reveals the development of geologic processes that complicates the understanding of the
paleontological record.

SANTAFÉ, J. V., CASANOVAS, M. L., SANZ, J. L. & CALZADA, S., 1982. Geología y paleontología (Dinosaurios) de las Capas Rojas
de Morella (Castellón, España). Diputación Provincial de Castellón y Diputación de Barcelona, Castellón y Barcelona (Eds.): 1-169.
SANTISTEBAN, C. DE, 2004. Ambiente sedimentario y ciclicidad estratigráfica de los materiales de la Formación "Arenas y arcillas del
Collado" (Cretácico inferior), en el sector de Alpuente (Valencia). Geogaceta, 35: 15-18.

Key words: Lower Cretaceous, Tethys, Eastern Iberia, eustatic cycles, dinosaur sites

86

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
NEW DINOSAUR FINDINGS FROM ARCILLAS DE MORELLA
FORMATION (SPAIN)

Andrés SANTOS-CUBEDO 1, 2, Carlos de SANTISTEBAN 3, Angel GALOBART 1

1 Institut Català de Paleontologia, c/ Escola Industrial 23, E-08201 Sabadell, Barcelona (ES),
angel.galobart@icp.cat
2 Grup Guix, c/ Santa Lucia 75, E-12540 Vila-real, Castellón (ES), andres.santos@icp.cat
3 Departament de Geologia, Universitat de València, Av. Dr. Moliner s/n, E-46100 Burjassot, València (ES),
Carlos.santisteban@uv.es

The Arcillas de Morella Formation (Aptian, Lower Cretaceous; eastern Iberian Chain, Spain) is widely
known by specialists because it has delivered an abundant and diverse collection of dinosaur remains (Galobart
et al. 2003; Ortega et al. 2006). Despite, only few skeletal fossils have been determined to specific or generic
level. Nowadays, the faunal dinosaur list from this formation is shown in Table 1 (references in Suñer et al.
2008):

Taxon
Site
Fossils
Thyreophora
Polacanthus
Cantera Mas de la Parreta
Postcranial
Ornithopoda
"Hypsilophodontidae"
Teuleria-Millán
Postcranial
cf. Hypsilophodon sp.
Royo y Gómez collection
Tooth
Iguanodon sp.
Povet de Sant Llàcer; Ana
Postcranial
I. bernissartensis
El Beltrán (Teuleria Azuvi); Cantera Mas de la
Postcranial,
Parreta; Mas Guimerà; Masia Eroles; Mas Romeu
Jaws, Teeth
Sauropoda
Titanosauriformes
Sant Antoni de la Vespa; El Canteret; Masia
Postcranial
Eroles; Cantera Mas de la Parreta
Theropoda
Theropoda indet.
Morella; Mas Romeu; El Beltrán; Ana
Teeth,
Postcranial
Allosauroidea indet.
Cantera Mas de la Parreta
Teeth,
Postcranial
?Carcharodontosauridae
El Beltrán
Tooth
Spinosauroidea
Vallibona
Postcranial
Spinosauridae
Cantera Mas de la Parreta
Postcranial
Baryonychinae indet.
Morella; Ana; Cantera Mas de la Parreta
Teeth,
Postcranial
Coelurosauria indet.
Cantera Mas de la Parreta
Teeth,
Postcranial

Table 1. Faunal list of the Arcillas de Morella Formation.

We report new fossil findings from the Arcillas de Morella Formation, including new
"Hypsilophodontidae" material from Mas del Dolço site, and new Iguanodontia and Titanosauriformes material
from Ana site. These new findings will help to understand the dinosaurs present in the Lower Cretaceous of Els
Ports (Castellón, Spain).

GALOBART, A., GAETE, R., SANTOS-CUBEDO, A., SUÑER, M., & VILA, B., 2003. New dinosaur sites in Catalonia and Valencia (J/K
boundary and Upper Cretaceous) and a short overwiew of Mesozoic dinosaur sites of Spain. The Palaeontology Newsletter. The
Palaeontological Association. Review Seminar on British Dinosaur, co-hosted by the Dinosaur Isle Museum and the Universtiy of
Portsmouth. Isle of Wight. Abstracts, 54:103-104.
ORTEGA, F., ESCASO, F., GASULLA, J.M., DANTAS, P., AND SANZ, J. L. 2006. Dinosaurios de la Península Ibérica. Estudios
Geológicos, 62(1): 219-240.
SUÑER, M., POZA, B., VILA, B., AND SANTOS-CUBEDO, A. 2008. Síntesis del registro fósil de dinosaurios en el Este de la Península
Ibérica. Palaeontologica Nova. SEPAZ, 8: 397-420.

Key words: Geological record, Arcillas de Morella Formation, Dinosaurs

87

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
ORGANIC MATTER CHARACTERIZATION AND ORGANIC
CARBON ISOTOPES RECORD IN THE EARLY CRETACEOUS
LACUSTRINE SETTING OF BERNISSART (BELGIUM)

Johann SCHNYDER 1, Jean DEJAX 2, Eddy KEPPENS 3, Thanh Thuy NGUYEN TU 4,
Paul SPAGNA 5, Armelle RIBOULLEAU 6, Johan YANS 7

1 UPMC Univ Paris 06, CNRS, UMR 7072, case postale 117, 4, pl. Jussieu, 75252 Paris Cedex 05, France,
johann.schnyder@upmc.fr
2 UMR MNHN-UPMC-CNRS 5143 "Paléobiodiversité et paléoenvironnements", case postale 38, 57 rue
Cuvier, 75231 Paris Cedex 05, France, dejax@mnhn.fr
3 Vrije Universiteit Brussel, Department of Geology, Pleinlaan 2, 1050 Brussels, Belgium,
ekeppens@vub.ac.be
4 UMR UPMC-CNRS-MNHN 5143 "Paléobiodiversité et paléoenvironnements", MNHN case postale 38, 57
rue Cuvier, 75231 Paris Cedex 05, France, Thanh-Thuy.Nguyen-Tu@snv.jussieu.fr
5 Faculté Polytechnique de Mons, rue de Houdain 9, 7000 Mons, Belgium, Paul.Spagna@fpms.ac.be
6 Université de Lille 1, UMR 8110 PBDS, bâtiment. SN5, cité scientifique, 59655 Villeneuve d'Ascq cedex,
France, armelle.riboulleau@univ-lille1.fr
7 Facultés Universitaires Notre-Dame de la Paix de Namur, 61 rue de Bruxelles, 5000 Namur, Belgium,
johan.yans@fundp.ac.be

We studied the sedimentary organic matter (OM) content of a 50-m-long core drilled in lacustrine
Wealden sediments (Middle Barremian-Early Aptian) from the dinosaur-bearing Bernissart setting (Belgium).
Bulk organic data (Rock-Eval, palynofacies, C/N ratios) and carbon isotope measurements on bulk OM and
selected isolated wood fragments were performed. The bulk OM distribution shows two organic cycles that are
stacked in a long-term cyclic pattern. This sequence has been divided into three units. Unit 1 at the base of the
core is characterized by rather low to moderate total organic carbon (TOC = 2.65 in wt % in average), relatively
low hydrogen index (HI = 11 to 203 mgHC/gTOC, average value: HI = 82 mgHC/gTOC) low proportions of
amorphous organic matter (AOM) and high proportions of vascular plant-derived OM in sieved palynofacies
slides. Units 2 and 3 are characterized by higher TOC (3.80 wt % in average), with an increasing and decreasing
trend, respectively, relatively higher HI (9 to 369 mg HC/gTOC, average value: HI = 159 mg HC/gTOC), and a
generally-dominating AOM content. These features are interpreted in terms of increasing in situ production and
preservation of OM through time within the freshwater masses. Carbon isotope curve of bulk OM is interpreted
as the result of two local patterns in sedimentation: (1) the varying contribution of in situ AOM vs. higher plant-
derived OM, and (2), the possible influence of enhanced productivity. The trend towards more negative values in
the carbon isotope records of wood is similar to that observed for bulk OM. Moreover, it is parallel to the one
previously recognized in coeval English Wealden facies using wood fragments. Therefore, this trend is regarded
as reflecting the changes of the carbon isotope ratios of atmospheric CO2 during the Middle Barremian-Early
Aptian.

Key words: Wealden facies, Barremian, Aptian, lake, palynofacies

88

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
MAMMALIAN FAUNA OF HYENA DEN CAVE (ALTAI, RUSSIA)

Natalia SERDYUK1, Sergei VASILIEV2

1 Paleontological Institute of RAS, 117997, Moscow, Profsoyuznaya str., 123, nataly@paleo.ru
2 Institute of archaeology and ethnography Siberian branch of RAS, 630090, Novosibirsk, pros. Akad.
Lavrentiev, 17, SVasiliev@archaeology.nsc.ru

The Hyena Den cave is situated in North-western Altai (Russia). It is located on the left bank of the Big
Tigirek River. The cave has a southwestern orientation. The entrance is at 115 m above the river level. The cave
was discovered by Russian scientist N.D. Ovodov in 1969. In 2008 the Institute of Archaeology and
Ethnography of the Siberian Branch and Paleontological Institute of the Russian Academy of Sciences started
new excavations of the site. The Hyena Den cave is a karst cavity in Early Silurian limestone. Cave consists of
two horizontal galleries. In 2008, a geological profile of untouched horizons was studied in the right gallery of
the cave. Pleistocene sequences of the Hyena Den cave have been divided into 6 levels. The base of the sequence
is formed by large rocky blocks, which do not represent the cave floor.
Many remains of large and small mammals have been discovered. The cave was named "Hyena Den"
because a great part of the bones was heavily broken, with tracks of gastric juice, and hyena's dairy teeth are
abundant. 3 800 bone fragments of large mammals have been unearthed; 17 % of them can be identified and
include the following taxa: Canis lupus, Vulpes vulpes, V, corsak, Mustela altaica, M. meles, Crocuta spelaea,
Panthera spelaea, Mammuthus primigenius, Equus ferus, Equus cf. hydruntinus, Equus sp., Coelodonta
antiquitatis, Cervus elaphus, Megaloceros giganteus, Bos (Pophagus) baicalensis, Bison priscus, Capra
sibirica, Ovis ammon, Ovis-Capra. Specific structure and bone abundance of large mammals does not change
from layer to layer.
More than six thousand small mammal bone fragments have been found in this cave. The structure of
small mammal fauna from Hyena Den cave is typical for Late Pleistocene fauna from Altai. According to small
mammals data, layers 1-2 are possibly Holocene in age and layers 3-6 Pleistocene. More than 30 small mammal
spcies have been recorded: Chiroptera gen indet, Asioscalops altaica, Sorex minutissimus, S. minutus, Crocidura
sp., Pteromys volans, Sciurus vulgaris, Eutamias sibiricus, Spermophilus sp., Marmota sp., Sicista sp., Allactaga
sp., Apodemus sp., Allocricetulus eversmanni, Cricetus cricetus, Ellobius sp., Miospalax myospalax,
Clethrionomys rufocanus, C. rutilus, C. glareolus, Cricetulus barabensis, Alticola strelzowi, A. macrotis,
Lagurus lagurus, Eolagurus sp., Arvicola terrestris, Stenocranius gregalis, Microtus oeconomus, M. agrestis, M.
arvalis, M. middendirfii, M. hyperboreus, Ochotona alpina-hyperborea, and O. sp.
Mammal faunas attest the presence, near Hyena Den cave of steppe landscapes and shrubbery associations
by Late Pleistocene time; forests were not extensive. The climate was mild without temperature leaps and
droughts.
The work was supported by the projects nos. 08-04-00483- of the Russian Foundation for Basic
Research.
Key words: Mammals, Late Pleistocene, environment, Altai, Russia

89

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
COMPARISON OF SEDIMENTOLOGICAL DATA IN TWO WEALDEN
FACIES SITES: THE BERNISSART NATURAL PIT AND THE
DANUBE-BOUCHON QUARRY OF HAUTRAGE (MONS BASIN,
BELGIUM). IMPLICATION ON THEIR GEODYNAMIC HISTORY

Paul SPAGNA 1, Christian DUPUIS 1, Johan YANS 2

1 UMons, GFA, 9 rue de Houdain, 7000 Mons, Belgium, Paul.Spagna@fpms.ac.be -
Christian.Dupuis@fpms.ac.be,
2 FUNDP UCL-Namur, Geology, 61 rue de Bruxelles, 5000 Namur, Belgium, Johan.yans@fundp.ac.be

This paper compares the wealden deposits of the Hautrage Danube-Bouchon quarry (Hautrage Clays
Formation) with those found in the recent boreholes drilled in the Bernissart natural clay pit (Sainte-Barbe Clays
Formation). These two formations have continental origin, and are middle Barremian to earliest Aptian in age
(Yans et al. 2006; Dejax et al. 2007). Moreover, they have been trapped and conserved till nowadays by
phenomena that can be compared in terms of mechanisms (anhydrites layers deep dissolution induced
subsidence) even if the scales of the subsiding area are different.

From a lithological point of view, Hautrage sediments (235 m-thick) are very heterogeneous, composed
of clays, silts, sands and even conglomeratic beds, with various contains of both sideritic and pyritic nodules, and
of pluricentimetrical to plurimetrical in size wood fragments. The environment associated to those deposits is
interpreted as a flooding plain crossed by numerous east-west channels (Spagna et al. 2008). The clayey
sediments (major part of the series) are related to lacustrine to swampy deposits that took place during high
rising water events when the overflow of the river banks leads to the sedimentation of the finest particles. Poor
maturation soils are frequent in the series (mainly at its bottom), indicating relatively long periods of exudation.
Sandy channels are mainly observed at the top of the series (southern front of the quarry), even if older (but
smaller) ones have also been described and studied in its lower part. Most of the pebble beds are localised on the
eastern part of the south front, probably associated to colluvial deposits, such as bank sliding, which could have
been induced by wildfire events (Spagna, in progress)

On the other hand, the 50 meters-thick of Sainte-Barbe Clays Formation recently cored (BER3) in the
"Cran aux Iguanodons" has been described as a very homogeneous series, composed of finely laminated dark
clays (Spagna & Van Itterbeeck, 2006) interpreted as varved sediments. The consistent evolution of both the
granulometric content and the organic matter properties (see Schnyder et al. this volume) allows the subdivision
of the wealden series in at least three zones.

At Bernissart the accommodation of the sediment in response to the subsidence progression is marked
by the creation of multiple plurimetrical blocks that moved separately during their downward progression. At
Hautrage, the bigger scale of the subsiding area leads to the formation of an extrado graben (described in
Vandycke & Spagna, this volume), localised just at the break of slope of the wealden strata. The graben
formation is suspected to affect the subsequent sedimentation dynamic by a quick induced creation of space,
settled by (lacustrine-like) dark clayey sediments (Spagna, in progress).

DEJAX J., PONS D. & YANS J. (2007). Palynology of the dinosaur-bearing Wealden facies sediments in the natural pit of Bernissart
(Belgium). Review of Palaeobotany and Palynology 144, 25-38 (IF 0.867).
SPAGNA P. & VAN ITTERBEECK J. (2006) Lithological description and granulometric study of the wealden facies in two borehole core
drilled in the "Cran aux Iguanodons de Bernissart" (N-W of the Mons Basin, Belgium). Poster - Geologica Belgica Meeting,
Liège.
SPAGNA P., DUPUIS C., & YANS, J. (2008) Sedimentology of the wealden clays in the Hautrage quarry. Memoirs of the Geological
Survey of Belgium n°55 pp. 35-44.
SPAGNA P. (in progress) Les facies wealdiens du Bassin de Mons (Belgique) : étude géodynamique, paléoenvironnementale et valorisation
industrielle. Phd Faculté Polytechnique de Mons, Mons.
YANS J., DEJAX J., PONS D., TAVERNE L. & BULTYNCK P. (2006). The Iguanodons of Bernissart are middle Barremian to earliest
Aptian in age. Bulletin Institut Sciences naturelles Belgique 76, 91-95.

Key words: Wealden, paleoenvironment, Hautrage, Bernissart, geodynamic

90

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE PROBLEMATIC FOSSIL BALEEN WHALES FROM BELGIUM

Mette E. STEEMAN

Biological Institute, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen Ø,
mesteeman@bio.ku.dk

The Royal Belgian Institute of Natural Sciences, Brussels, houses a large historical collection of Middle
Miocene to Pliocene marine mammals, collected during expansions of the fortifications of Antwerp during the
middle of the 19th century. Based on the vast amount of material, P.-J. Van Beneden published beautifully
illustrated descriptions of, amongst others, twenty species of baleen whales that do not belong to any of the
recent families. They became the foundation of the future taxonomy and understanding of later described extinct
mysticetes and they include widely used genera such as Plesiocetus and Mesocetus. To this day, these species
make up about one third of all described baleen-bearing mysticetes not belonging to recent families. However,
Van Benedens mysticete species have also proved to be problematic. Most of them are based on several syntypes
consisting of single isolated elements from various parts of the skeleton, and they are likely to have originated
from more than one biological species. Because different syntypes subsequently have been used for comparison
for later described material, species that clearly are not closely related have been referred to the same genus (e.g.
Mesocetus). The need for a revision and establishment of lectotype material is therefore apparent. Cetacean ear-
bones are recognised as having good diagnostic value, with the periotic slightly more so than the bulla.
Fortunately, the periotic consists mainly of very compact bone and usually preserves well. Most of Van
Beneden's species contain a periotic among the syntype material. Using the periotic, it is therefore possible to
establish comparable and diagnostic lectotypes for several of Van Benedens species. It should be emphasized
that it is not recommended to establish new species based on such limited material, but by using the periotic as
lectotypes here, it is possible to define these well-established and widely cited genera. Comparing these periotics
to those of the better preserved, later described species belonging to a contemporary East Atlantic fauna shows
that the diversity of these mysticete faunas were comparable.

Key words: Van Beneden, Mysticeti, Taxonomy, Miocene

Figure: Illustration published by P.-J. Van Beneden (1886) of a partial basicranium including a periotic and
bulla from Idiocetus longifrons (one of the most complete syntypes).

91

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
QUANTIFYING GROWTH RATES IN ISLAND DWARF SAUROPODS

Koen STEIN, Martin SANDER

Steinmann Institute, Division Palaeontology, Nussallee 8, 53115 Bonn, Germany
koen.stein@uni-bonn.de

Europasaurus holgeri and Magyarosaurus dacus are both of small stature, and the only unequivocal
island dwarf sauropods known so far. A preliminary histological description of Europasaurus long bones proved
the largest individual was fully grown at about 6m length. The historically debated island dwarf status of
Magyarosaurus dacus (Nopcsa 1914; Jianu and Weishampel, 1999; Leloeuff, 2005) has recently been confirmed
by long bone histological analysis (Stein et al. in Prep.). However, secondary remodelling in Magyarosaurus
was so advanced no statements could be made on its growth rate other than it was reduced.
Europasaurus long bones preserve the primary growth record, with cortical growth marks. Minimal age
estimates of the studied specimens were obtained by counting and backcalculating cortical growth marks. In total
5 individuals, estimated at 1, 3, 6, 7 and 17 years minimal age, were used to construct a growth curve. The
largest individual displays an external fundamental system, and was estimated at 17 years minimal age. Using
longbone circumference, it was estimated to have a mass of 690 kg. Estimates of mass for the other individuals
were obtained using developmental mass extrapolation (Erickson et al. 2000). All individuals plotted on a
smooth growth curve.
Growth curve data show that Europasaurus had a maximal average growth rate of 74 kg/year (± 217
g/day). On Case's (1978) chart with maximum growth rate versus adult body mass, Europasaurus plots on the
general dinosaur regression line. However, it has moved away from the sauropods with extremely high growth
rates, towards the prosauropod condition. This means it decreased its growth rate with a factor 10. This suggests
that sauropod growth rates are highly flexible, and under the right conditions can rapidly be increased as well as
decreased, analogous to mammals.

CASE, T. R., 1978. Speculations on the growth rate and reproduction of some dinosaurs. Paleobiology, 4: 320-328.
ERICKSON, G. M., AND TUMANOVA, T.A., 2000. Growth curve of Psittacosaurus mongoliensis Osborn (Ceratopsia: Psittacosauridae)
inferred from long bone histology. Zoological Journal of the Linnean Society, 130: 551-566.
JIANU, C. M. AND WEISHAMPEL, D.B., 1999. The smallest of the largest: a new look at possible dwarfing in sauropod dinosaurs.
Geologie en Mijnbouw, 78: 335-343.
LE LOEUFF, J., 2005. Romanian Late cretaceous dinosaurs: big dwarfs or small giants? Historical Biology, 17: 15-17.
NOPCSA, F. von, 1914. Über das Vorkommen der Dinosaurier in Siebenbürgen. Verhandlungen der Zoologisch-Botanischen Gesellschaft,
54: 12-14.

Key words: sauropods, growth, Europe, islands, dwarfism

92

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE AFRICAN COUSINS OF THE EUROPEAN IGUANODONS

Philippe TAQUET

Muséum National d'Histoire Naturelle, 8 rue Buffon, Paris 75005, France, taquet@mnhn.fr

The discovery in Africa in 1966 of the skeletons of two new Iguanodons in the sediments of the
Gadoufaoua locality (Niger), dating from the Aptian (Lower Cretaceous), has allowed us to describe two new
genera: the first one, Ouranosaurus nigeriensis is a gracile Iguanodontid, with a facultative bipedality, with
bumped nasals on the skull and long neural spines on the dorsal vertebrae. The second one, Lurdusaurus
arenatus, is a heavy, quadrupedal Iguanodontid, with very short and robust limbs and a hippopotamus-like body.
These two African Iguanodons are very probably descendants from the European Iguanodons, as Iguanodon
atherfieldensis and Iguanodon bernissartensis; their ancestors were migrating from Europe to Africa during the
Lower Cretaceous.
Considering the Aptian datation of the Gadoufaoua locality and the morphologic evolution of the Nigerian
Iguanodons, it was suggested in 1975 a Barremian age for the Belgian Iguanodons; this age was confirmed
recently by a palynological dating of the Wealden sediments of Bernissart (Mons Basin, Belgium).

Key words: Iguanodons, Aptian, Barremian, Niger, Belgium

93

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
FIRST MAASTRICHTIAN VERTEBRATE ASSEMBLAGE FROM
PROVENCE (VITROLLES-LA-PLAINE, FRANCE)

Xavier VALENTIN 1, Pascal GODEFROIT 2, Rodolphe TABUCE 3,
Monique VIANEY-LIAUD 3, Géraldine GARCIA 1, Wenhao WU 2

1IPHEP, Université de Poitiers, 40 Avenue du Recteur Pineau, 86022 Poitiers cedex, France,
geraldine.garcia@univ-poitiers.fr
2 Department of Palaeontology, Royal Belgium Institute of Natural Sciences, rue Vautier 29, 1000 Brussels,
Belgium, Pascal.Godefroit@naturalsciences.be
3 ISEM, Université Montpellier II, place Eugène Bataillon, 34095 Montpellier cedex 05, France,
rodolphe.tabuce@univ-montp2.fr

A new Maastrichtian locality from Provence (southwestern France) has yielded a diversified vertebrate
fauna, including a zhelesthid mammal, Valentinella vitrollense, (Tabuce et al., 2004) associated with several
groups of reptiles (Squamates, Chelonians, Crocodilians) and dinosaur families (Abelisauridae,
Dromaeosauridae, Titanosauridae, Rhabdodontidae, Hadrosauridae). It is the first noticeable report of the
occurrence of hadrosaurids in Provence. The morphology of the dentary and dentary teeth suggests that the
hadrosaurid material belongs to a basal form. It is clearly different from hadrosaurid material previously
described in Europe. However, the material discovered so far is too fragmentary to erect a new taxon. This new
site offers new perspectives on the diversity and evolution of the European vertebrate ecosystems during the
Maastrichtian. Indeed, the association of hadrosaurids with titanosaurids and Rhabdodon is unusual in the latest
Cretaceous of southern France. It questions the hypothesis of the replacement of a Early Maastrichtian fauna
dominated by titanosaurid sauropods by a Late Maastrichtian assemblage dominated by hadrosaurs during the
Maastrichtian in western Europe (e.g. Le Loeuff et al., 1994).

LE LOEUFF, J., BUFFETAUT, E., & MARTIN, M., 1994. The last stages of dinosaur faunal history in Europe : a succession of
Maastrichtian dinosaur assemblages from the Corbières (southern France). Geological Magazine, 131: 625-630.
TABUCE, R. et al., 2004. A eutherian mammal in the latest Cretaceous of Vitrolles, southern France. Acta Paleontologica Polonica, 49:
347-356.

Key words: vertebrates, Maastrichtian, Provence, hadrosaurids, dinosaur assemblages

94

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
CRUSTAL TECTONIC CONTROL OF THE EARLY CRETACEOUS
DEPOSITS IN THE MONS BASIN (BELGIUM)

Sara VANDYCKE 1, Paul SPAGNA 2

1 Géologie Fondamentale et Appliquée, UMons, Research Associate FNRS, Sara.Vandycke@fpms.ac.be
2 Géologie Fondamentale et Appliquée, UMons, Paul.Spagna@fpms.ac.be

In the Mesocenozoic structural geological network of the NW Europe, the Mons Basin is located on a
major inversion crustal zone. Active tectonics during the Cretaceous induced extensional and strike-slip system
well recorded in the sediments of the Mons Basin in terms of brittle structures. The deformation analysis of early
Cretaceous Wealden sediments helps to understand early stages of the Cretaceous basin development. This
present analysis has been done mainly in the Danube-Bouchon at Hautrage, after excavating campaigns. This
study also details the relationship between regional tectonic activity and the deposit process.
In the Mons Basin, the outcropping Wealden facies deposits are located on the north edge of the Mons
Basin, where they appear in plurikilometric-sized pockets (Hautrage, Baudour, Thieu, ..), or in fillings of smaller
natural pits (Bernissart). The investigated Hautrage Clays Formation is middle Barremian to earliest Aptian in
age. It is composed of continental clays, silts and sands containing lignite remains, nodular pyrite and siderite
nodules in variable proportions. The depositional environment of Hautrage, in the Danube-Bouchon Quarry is
interpreted as a flooding plain crossed by E-W numerous channels.
Different types of tectonic structures and deformations are observed. First, spectacular hydraulic faulting
with hydroplastic contrasted injection of white-pink clays injection in the black sediments was discovered.
Displacement has been recognized (around 10 meters) but surfaces of fault are not striated. Those hydroplastic
injection structures are oriented around E-W. Some of them are indeed deformed by compaction. Secondly,
multi-stages of normal faulting are structured in a major E-W graben structure. Centimetric to pluri-metric
displacements are noted, especially in the channel basal bed. This normal faulting coincides with the beginning
of the Mons Basin. Localised on the break of slope of the Wealden strata, the opening of the graben is
interpreted as a response (accommodation) to a southward subsidence front movement. On the other hand,
dextral NE-SW strike-slip faults have been observed. These structures cut the E-W graben. Lateral fault
displacement has been proved by sub-horizontal slicken-sides but also by lateral displacement of the main beds
formations observed during the progressive excavation. Oblique jointing has also been observed. This strike-slip
system is related to regional tectonics. At least, well organized NW-SE extensional joints are nicely developed in
the black clays beds. This system of jointing can be related to NE-SW extensional tectonics well known in the
Mons Basin until the Upper Cretaceous but also in the whole NW Europe related to the opening of the Lower
Rhin Graben.
To conclude, tectonic analysis in Wealden sediments, not necessarily well competent in terms of rocks
mechanics, helps anyway to precise the geodynamic environment during the Early Cretaceous.

Key words: Wealden, tectonics, Mons Basin, clays, faulting

95

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
ORIGINE DE L'HOMME: LES NOUVELLES DU TCHAD

Patrick VIGNAUD 1, MPFT 2

1IPHEP - CNRS UMR 6046 - UFR SFA Université de Poitiers 40 av. du Recteur Pineau 86022 POITIERS
cedex
2La Mission Paléoanthropologique Franco-Tchadienne est une structure de collaboration scientifique entre
l'Université de Poitiers et le CNRS (France), l'Université de N'Djaména et le CNAR (Tchad)

L'Afrique, continent-clé pour l'origine et l'évolution des Hominidés, offre en plusieurs régions de
longues séquences sédimentaires (Mio-Plio-Pléistocène) où sont enregistrés les événements géologiques et
climatiques qui canalisent l'évolution des paysages et des systèmes biologiques. Dans ce contexte, le Bassin
intracratonique du Lac Tchad, aujourd'hui en grande partie désertique et soumis à une intense érosion éolienne,
renferme de très nombreux sites fossilifères datés du Miocène sup. à l'Holocène. Il constitue une nouvelle
fenêtre, ouverte sur la connaissance de l'origine et de l'évolution des Hominidés anciens.
En 1995, au nord du Tchad, la Mission Paléoanthropologique Franco-Tchadienne (MPFT) met au jour
"Abel", de son nom scientifique Australopithecus bahrelghazali Brunet et al 1996, le premier Australopithèque
connu à l'Ouest de la Rift Valley. Il est associé à une faune qui indique un âge biochronologique de 3 à 3,5 Ma.
En 2001, la découverte de Toumaï, le plus ancien Hominidé connu, dans des sédiments datés de 7 Ma, conduit à
revoir de manière drastique les conceptions de l'histoire des premières phases du rameau humain.
Ce crâne complet mais déformé et fracturé au cours de l'enfouissement, a été scanné, "déconstruit" puis
reconstruit en réalité virtuelle 3D. Il est alors possible d'intégrer les caractères morphométriques dans l'étude du
crâne de Sahelanthropus tchadensis et de ses relations phylogéniques. L'ensemble des caractères anatomiques et
morphométriques du crâne montre que Sahelanthropus tchadensis appartient bien au rameau humain. Très
probablement bipède, il est proche de la divergence avec le groupe des chimpanzés. De ce fait, cette découverte
repousse à plus de 7 Ma l'âge de cette dichotomie classiquement proposée par les phylogénies moléculaires entre
5 et 6 Ma...
Dans le Mio-Pliocène, plus de 500 sites paléontologiques ont maintenant été mis au jour sur 4 secteurs
fossilifères de composition faunique et d'âge différents entre 3 et 7 Ma. Plus de 15 000 fossiles ont été collectés,
déterminés et inventoriés. Les faunes et les flores sont étudiées en tant que composantes biologiques des
écosystèmes. L'étude exhaustive de la faune (taxinomie, phylogénie, analyses paléoécologiques, etc.) est en
cours mais elle permet d'ores et déjà de dresser des schémas environnementaux dans lesquels ont évolué les
Hominidés d'Afrique Centrale depuis le Miocène supérieur.
Dans le Miocène supérieur (secteur de Toros-Menalla, ca. 7 Ma), la faune dessine un paysage mosaïque
allant de milieux aquatiques à humides (Poissons, Crocodiliens, Tortues, Hippopotamidés, Anthracothéridés,
etc.), avec forêts claires et savanes arborées (Proboscidiens, Bovidés, Giraffidés, Primates, etc.) à des milieux
plus ouverts de prairies à graminées (Lagomorphes, Equidés, Bovidés, etc.).
Mis à part de nombreux restes de légumineuses lianescentes, les macro-restes de végétaux sont absents
des sites tchadiens ; les phytolithes et diatomites sont actuellement en cours d'étude. Le couvert végétal est donc
approché par des méthodes indirectes (biogéochimie isotopique et stries d'usure dentaire des grands mammifères
herbivores). Après caractérisation précise de la diagenèse, l'analyse isotopique de l'émail dentaire (13C et 18O sur
la carbonate hydroxylapatite) a permis notamment de montrer une certaine ouverture des milieux entre 3 et 7 Ma
en Afrique Centrale (résultats en cours de publication). L'étude des usures dentaires des herbivores est rendue
difficile par la rareté des structures conservées dans cette région désertique où l'érosion éolienne est très intense.
Les données de la paléontologie sont confortées par le contexte sédimentologiques. Les séries mio-pliocènes
étudiées (entre 3 et 7 Ma) correspondent à une succession de phases humides (lacs), semi-arides (fleuves-
savanes), voire arides (désert) dans les niveaux les plus anciens. Diatomites et argiles marquent l'installation
d'épisodes lacustres francs qui transgressent à grande échelle sur les faciès fluviatiles.
Enfin, la position géographique du Bassin du Lac Tchad est importante pour la compréhension des
différents échanges fauniques au cours du Mio-Pliocène africain. Les premiers résultats portant sur certains
groupes taxonomiques (Poissons, Crocodiliens, Proboscidiens, Hippopotamidés, Carnivores, etc.) montrent le
rôle déterminant de l'Afrique Centrale dans la mise en place des échanges fauniques intermittents avec l'Afrique
de l'Est, l'Afrique du Nord et l'Eurasie.

Key words : Chad, Mio-Pliocene, Hominidae, palaeoenvironments

96

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
DIVERSITY AND PALEOGEOGRAPHIC DISTRIBUTION OF EARLY
JURASSIC PLESIOSAURS

Peggy VINCENT 1, Guillaume SUAN 2

1 UMR 5143, Paléobiodiversité et Paléoenvironnements, Département Histoire de la Terre, Muséum national
d'histoire naturelle, CP 38-75231 Paris cedex 05, France
2 UMR 5125, PaléoEnvironnements et PaléobioSphère, Bâtiment Géode, Campus de la Doua, Université Lyon
1, 69622 Villeurbanne, France

Early Jurassic plesiosaurs, a group of extinct marine reptiles, were one of the first groups to be
described in the history of vertebrate paleontology. Nevertheless, the paleogeographic distribution and the
taxonomic diversity of these forms are still unclear, particularly because most descriptions and taxonomic
attributions were realized during the mid 19th to early 20th century. Here we investigate the paleodiversity and
paleogeographic distribution of Early Jurassic plesiosaurs using an extensive taxonomic and anatomical revision
of most known Early Jurassic specimens. We also present an examination of the biostratigraphic and
sedimentological framework of deposits in which these specimens were discovered, in order to decipher whether
their fossil record reflects primary paleobiological trends or taphonomic/discovery biases. Early Jurassic
Plesiosaur diversity appears to reach its maximum during the Toarcian (falciferum-bifrons ammonite zones).
Nevertheless, the inclusion of ghost lineages (i.d. lineages which presence is suggested by the phylogenetic
relationships, but not present in the fossil record) into the diversity curves indicates that this pattern mostly
reflects the consequence of discovery and taphonomical biases rather than primary biodiversity trends. Indeed,
most strata where numerous plesiosaurs species were discovered correspond to sediments that were both
deposited under poorly-oxygenated conditions and exploited at least in a semi-industrial way during the 1800's-
1950's. The Lower Jurassic fossiliferous localities that yielded identifiable plesiosaur species are only found in
Western Europe (England, Germany, and France). In Europe, the Toarcian stage is the only interval where more
than one fossiliferous locality is known (The Hettangian, Sinemurian and Pliensbachian stages being each
represented by only one locality where specimens are identifiable at the species level). The different Toarcian
fossiliferous sites of Europe do not bear any single common taxon, suggesting a high degree of endemism in
Early Jurassic plesiosaurs. Nevertheless, these sites are fundamentally diachronous at the ammonite zone level;
this absence of shared taxa might hence reflect temporal changes rather than paleogeographic trends. Further
data are required to determine whether if this pattern is a consequence of truly limited paleobiogeographic ranges
or the result of high rates of turnover. In addition, future fossil discoveries and refinements of the phylogenetic
relationships are required to precise the evolution of this diversity at a higher stratigraphic resolution, and hence
determine how plesiosaurs responded to severe environmental change that punctuated this period (i.e. Early
Hettangian and Early Toarcian mass extinction events).

Key words: Early Jurassic, Plesiosauria, Europe, biostratigraphy, palaeogeography

97

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
THE TERRESTRIAL LOWER CRETACEOUS OF GERMANY
("WEALDEN")

Volker WILDE 1, Walter RIEGEL 1,2

1 Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Senckenberganlage 25, D - 60325 Frankfurt
am Main, Germany, Volker.Wilde@senckenberg.de
2 Geowissenschaftliches Zentrum der Universität Göttingen, Abteilung Geobiologie, Goldschmidtstr. 3, D -
37077 Göttingen, Germany, wriegel@gwdg.de

The term "Wealden" has been applied to more or less nonmarine sediments of Lower Cretaceous age,
especially in Europe but also in other parts of the world. Except for the "type"-Wealden in Southern England, the
term "Wealden-facies" should be preferred. Wealden-facies sediments have been described especially from
Western Europe (England, Spain, France, Belgium, The Netherlands), the northern part of Germany, and
Southern Scandinavia. By far most of the Northwest German Wealden-facies is representing the Berriasian to
early Valanginian filling of a large restricted basin extending from the area of the Dutch-German border in the
West to an area North of Braunschweig. To the South it was bounded by the Rhenish Massif, and to the North by
the Pompecki-Swell or Rise. Intermittent connections to the marine realm have been suggested especially to the
North and to the West. The Northwest German Wealden-facies is known from extensive outcrops along the
southern margin of the basin representing a variety of fluvial to lacustrine sediments including clays, sandstones,
and coals. The major barrier sandstones are still of economic interest and became recently famous for their
frequent and well preserved dinosaur trackways. The central and northern part of the basin is deeply concealed
by Quarternary sediments and especially known from oil exploration wells of the middle of the last century. It is
occupied by bituminous clays with intercalations of monotypic shell beds and marginal clastic intercalations.

More or less isolated occurrences of Wealden-facies sediments in the eastern parts of Germany and on
the Pompecki-Swell are only known from deep wells and may be of slightly different age, somehow between
Berriasian and Barremian. Another type of Wealden-facies sediments is today restricted to isolated karst
structures on carbonate platforms in the northern part of the Rhenish massif. They are representing a
karstification "event" or phase leaving structures now filled with Barremian-age clays and sands rich in fusain
and sometimes yielding dinosaur bones, especially Iguanodon (the same phase or "event" is represented by the
sediments at Bernissart or Hautrage in Belgium).

Key words: Wealden, facies, Germany, Lower Cretaceous

98

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
A SAUROPOD DINOSAUR FROM THE CRETACEOUS OF JIUTAI,
JILIN, CHINA

Wen-hao WU 1, Zhi-ming DONG 1, 2, Yue-wu SUN 1, Chun-tian LI 1, Tao LI 3

1 Research Center of Palaeontology & Stratigraphy, Jilin University, Changchun 130026, Jilin, China
2 Institute of Vertebrate Paleontology and Paleoanthropology, CAS, Beijing 100044, China
3 Museum of Jilin University, Jilin University, Changchun 130026, Jilin, China

New basal titanosauriforms have recently been discovered all around the world and especially in China.
Here we describe a titanosauriform dinosaur, Jiutaisaurus xidiensis (Wu et al., 2006) from the Cretaceous
Quantou Formation (`middle' Cretaceous) exposed in Xidi village of Jiutai (Jilin Province, Northeastern China).
The holotype consists of a series of 18 articulated anterior and middle caudal vertebrae, with 13 associated
chevrons. The caudal bones are mainly composed of spongy tissue, made up of tiny cells. All the caudal
vertebrae are amphicoelous with the posterior face slightly more concave than the anterior. The caudal centra are
slightly constricted at their midlength. The neural arches are situated on the anterior half of the centra. The
diameters of the centra and the heights of the neural spins decrease progressively caudally. There are incomplete
transverse processes preserved in the anterior caudal centra, and the chevron surfaces at the ventral surface of the
centrum are well developed, the articular surfaces are circular in the anterior caudal centra and sub-circular in the
middle caudal centra. There is a longitudinal prominence on the lateral surface of the middle centra. The
chevrons are not bridged at proximal ends and not unforked at their distal ends, and the haemal canal length is
more than half of chevron length.
Jiutaisaurus displays several key titanosauriform features, such as neural arches situated on the anterior
half of the centra, simple and undivided chevron blades, blades of middle chevron curved backward and
downward. Here Jiutaisaurus is regarded as a basal titanosauriform dinosaurs because it lacks the procoelous
anterior and middle caudals, regarded synapomorphic for more derived titanosaurines. But it also possesses two
key features for derived titanosaurines: spongy bone texture of caudal centra and the large heamal canal.
Recently, several analyses have focused on the Titanosauria phylogenetic relationships, however,
interrelationships among basal titanosauriforms are still not well understood. Unusual features in Jiutaisaurus
also indicate that titanosauriform evolution may be more complicated than was previously suspected.

Key words: Cretaceous, Dinosauria, Sauropoda, Titanosauriformes

99

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
INTEGRATED RESULTS OF THE NEW MATERIAL FROM THE 2002-
2003 BOREHOLES IN BERNISSART (MONS BASIN, BELGIUM)

Johan YANS 1, Bernard ANDREU 2, Jean-Marc BAELE 3, Colette CORNET 1, Armand
DE RICQLES 4, Jean DEJAX 5, Christian DUPUIS 3, Bruno GALBRUN 4, Thomas
GERARDS 6, Philippe GERRIENNE 6, Pascal GODEFROIT 7, Bernard GOMEZ 8,
René GOSSELIN 9, Thierry LEDUC 7, Gilles PETIT 4, Denise PONS 4, Alain PREAT 10,
Francis ROBASZYNSKI 3, Johann SCHNYDER 4, Thierry SMITH 7, Paul SPAGNA 3,
Etienne STEURBAUT 7, Louis TAVERNE 7, Jean-Pierre TSHIBANGU 11, Jimmy VAN
ITTERBEECK 7, Sara VANDYCKE 3, Céline VANNESTE 12

1 FUNDP Geology, UCL-Namur, Belgium, Johan.Yans@fundp.ac.be, Colette.Cornet@fundp.ac.be
2 Université Paul Sabatier, Toulouse, France, andreu@lmtg.obs-mip.fr
3 UMons, GFA, Mons, Belgium, Paul.Spagna@fpms.ac.be, Christian.Dupuis@fpms.ac.be
Sara.Vandycke@fpms.ac.be, Jean-Marc.Baele@fpms.ac.be, Francis.Robaszynski@fpms.ac.be
4 Université Pierre et Marie Curie-Paris 6, Paris, France, johann.schnyder@upmc.fr,
bruno.galbrun@upmc.fr, dpons@snv.jussieu.fr, armand.de ricqles@upmc.fr, Gilles.Petit@snv.jussieu.fr
5 MNHN, Département "Histoire de la Terre", Paris, France, dejax@mnhn.fr
6 ULg, PPM, Département de Géologie, Liège, Belgium, p.gerrienne@ulg.ac.be, tgerards@hotmail.com
7 IRSNB, Brussels, Belgium, Pascal.Godefroit@naturalsciences.be, Thierry.Smith@naturalsciences.be
Etienne.Steurbaut@naturalsciences.be, Thierry.Leduc@naturalsciences.be
8 Université Lyon 1 (Claude Bernard), Villeurbanne, France, bernard.gomez@univ-lyon1.fr
9 ASBL Patrimoine Iguanodons, Centre administratif du Préau, rue du Fraity 76, Bernissart
10 Université Libre de Bruxelles, DSTE, Bruxelles, Belgique, apreat@ulb.ac.be
11 UMons, Service Génie Minier, 53 rue du Joncquois, 7000 Mons, Katshidikaya.Tshibangu@fpms.ac.be
12 Région Wallonne, Cartographie géologique, Namur, Belgique, C.Vanneste@mrw.wallonie.be

Three new boreholes have been drilled within and around the natural pit ("Cran") of Bernissart in 2002-
2003. They provided exceptional material used for a multidisciplinary research to improve our knowledge of the
Iguanodons-bearing Wealden facies. The BER3 borehole reached 349.95 meters of Thanetian, Late Cretaceous,
Early Cretaceous and Westphalian sediments. Below 8 meters of green sandy clays it cuts 67 meters of Late
Cretaceous chalks and marls with local glauconites and cherts. The Albian and Cenomanian "meule" sediments
lie between 75 and 265 meters. The Wealden facies trapped in the "Cran aux iguanodons" of Bernissart are
recognised between 265 and 315 meters. The latter are laminated dark pyritic clays with mm-thick brown and
white silty levels. The borehole ends within a breccia with Carboniferous blocks between 315 and 349.95 meters.
Several topics were investigated such as palynolgy and determination of woody and plant-mesofossils fragments,
to detail the age and paleoenvironments of the Iguanodons. The formation processes of the "Cran" were
documented by multidisciplinary approach, i.e sedimentology of the lacustrine Wealden facies (including clay
mineralogy, granulometry and magnetic susceptibility), characterization of the organic matter with Rock-eval,
palynofacies, soluble alkane content, carbon isotope and structural analysis. Paleontological content was also
studied by palaeohistology and diagenesis of the bone fragments, characterization of amber, preparations for
diatoms and ostracods analyses (barren). Moreover, Wealden facies from IRSNB collection (historical searches
of 1878-1881) and other sites in the Mons basin (Hautrage, Thieu, Baudour) were also investigated, notably for
their content of numerous fishes and coprolites. In Hautrage and Baudour, we note the discovery of teeth of
freshwater sharks and a shinbone of a juvenile sauropod. In Thieu the occurrence of dinoflagellate cysts suggests
the marine influence for the Wealden facies in the Eastern part of the Mons basin.

Key words: Bernissart, natural pit, Wealden facies, Mons basin, boreholes

100

Darwin-Bernissart meeting, Brussels, February 9-13, 2009
CARBON ISOTOPES ON WOODY MATERIAL FROM WEALDEN
FACIES OF HAUTRAGE (MONS BASIN, BELGIUM)

Johan YANS 1, Jean DEJAX 2, Thomas GERARDS 3, Philippe GERRIENNE 3, Paul
SPAGNA 4, Eddy KEPPENS 5

1 FUNDP UCL-Namur, Geology, rue de Bruxelles 61, 5000 Namur, Belgium, Johan.Yans@fundp.ac.be
2 Museum d'Histoire Naturelle de Paris, Rue Buffon, 8, 75005 Paris, France, dejax@mnhn.fr
3 ULg, Laboratoire de Paléobotanique, Allée du 6 Août, Sart Tilman, 4000 Liège 1, Belgium,
p.gerrienne@ulg.ac.be, tgerards@hotmail.com
4 Faculté Polytechnique de Mons, GFA, rue de Houdain 9, 7000 Mons, Belgium, Paul.Spagna@fpms.ac.be
5 VUB, Geology, Pleinlaan 2, 1050 Brussels, Belgium, ekeppens@vub.ac.be

Carbon isotope ratios are a powerful tool reflecting geological events through their connection to the
global carbon cycle. They are used as a potential chemostratigraphic tool for geological successions, including
the main boundaries. Correlations are found through matching carbon isotope profiles in different sections from
pelagic to continental environments. The reliability of carbon isotopes has been demonstrated for bulk rocks,
pedogenic nodules, calcitic shells, dispersed organic carbon, but no systematic study has been done until now on
fossil woods. A thick succession of the "Wealden" facies sediments of the Mons Basin (Belgium) crops out in
the Danube-Bouchon quarry at Hautrage (Hautrage Clays Formation). This 235-m-thick succession encompasses
mainly dark to grey clays and sands, rich in organic matter and fossil woods, deposited in an alluvial plain. The
palynomorphs, including the primitive pollen of angiosperms, suggest a Middle Barremian to Earliest Aptian
age. We measured the carbon isotope ratios (110 levels) of 1) dispersed organic carbon (DOC), and 2) one fossil
wood (WOOD) collected in the same geological level. We aim to compare the two signals (DOC and WOOD) to
have a direct comparison in a continental stratigraphic section. The 13C of DOC and WOOD from the same
geological level are significantly different (average difference about 0.9). The 13C of the fossil-woods are
usually more positive than the 13C of DOC. The variability within the WOOD 13C is more important than in
the DOC 13C. Four main hypotheses may explain these results. Firstly, the 13C on DOC averages the isotopic
signal of different compounds and tissues (leaves, wood, and seeds) as they become mixed and dispersed. The
13C on WOOD reflects the carbon isotope ratio of a small part of one single tree. It is known that a single tree is
a complex system where we can observe 13C variations over a range of 4. Secondly, there is a strong 13C
variability between different species of plants. In Hautrage, several families were collected. Thirdly, diagenesis
can affect the 13C of the DOC and the various kinds of woods in a different way. Fourthly, the fossil woods may
be reworked several times from more ancient geological levels, especially in the alluvial plain environment of
Hautrage where thick levels of fluviatile coarse sands are observed. However, in the whole succession, both
DOC and WOOD 13C profiles show a positive trend. This can be due to several causes, including global pCO2
variations and/or changes in bioproduction and/or other environmental conditions in the alluvial plain. This
shows that carbon isotope profiles on wood material can be matched to carbon isotope profiles on DOC,
suggesting that fossil woods are a powerful chemostratigraphic tool when sufficient samples are measured in the
studied succession. Whenever possible a control of the wood taxon is recommended.

Key words: Wealden facies, Mons Basin, carbon isotopes, wood, dispersed organic carbon (DOC)

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FAUNA, FLORA, AND PALEOENVIRONMENT OF THE UPPER
DEVONIAN TETRAPOD-BEARING LOCALITY OF STRUD,
BELGIUM

Gaël CLEMENT 1, Cyrille PRESTIANNI 2

1 Département Histoire de la Terre, UMR 5143 du CNRS, Muséum national d'Histoire naturelle, 57 rue
Cuvier, 75231 Paris cedex 05, France, gclement@mnhn.fr
2 Unité de Paléobotanique - Paléopalynologie - Micropaléontologie (PPM), Département de Géologie,
Université de Liège, Allée du 6 Août, Bâtiment B-18, 4000 Liège 1, Belgium, cyrille.prestianni@ulg.ac.be

1. Introduction

Famennian outcrops of Belgium are located in the Namur and Dinant Synclinoria, the Vesdre Nappe
and the Theux Window. According to Bultynck & Dejonghe (2002), the Famennian stage, which is mainly
composed of siliciclastic rocks, corresponds to a regressive megasequence with oscillating movements due to
short terms transgressive pulses. Two main groups are distinguished, namely the Famenne and the Condroz
Groups. Depositional environments comprise juxtaposed continental, restricted marine and shallow marine
environments, including alluvial, lagoonal, evaporitic, tidal, sand barrier, and fore-barrier settings. A new
transgressive system, consisting of interbedded siliciclastic and carbonate rocks, overlay the Condroz Group.
Most of the quarries, used since the 19th century as building sandstone, are now abandoned but still
accessible and few of them are still intensively exploited.
The recent finding of a Famennian Ichthyostega-like tetrapod in Belgium (Clément et al., 2004) in the
Paleontology Collection of the Liège University was unexpected and, given the extreme rarity of Devonian
tetrapods worldwide, an important discovery.
On the contrary to almost all Devonian tetrapod-bearing localities of the world, the Famennian `Old
Red Sandstone' of Belgium is very well known. The Famenne region (Ardenne, southeastern Belgium) is the
stratotype area for the Famennian stage. Sedimentology, stratigraphy, tectonics, as well as micropaleontology of
the Famennian of Belgium have been studied in great detail for more than a century. However, large organismal
remains were considered too rare and useless to be studied in detail. As a result, most vertebrate reports date
back to the nineteenth century and the early twentieth century.
Recent field works were focused on the rediscovery of the tetrapod-bearing locality. "Strud" was the
only word engraved on the matrix of the tetrapod lower jaw and was the only clue to find the original outcrop. It
rapidly appeared that an important plant and vertebrate material from Strud collected at the end of the 19th
century is mainly housed in the IRSNB and Liège collections (completed by minor collections in Mons and
Louvain-La-Neuve Universities). This quarry was first investigated by Hock (1878) and plant and vertebrate
fossil material from Strud was later published by Lohest (1888), Leriche (1931) and Stockmans (1948). In
collaboration with geologists of the Liège University, the one-century abandoned quarry of Strud was
rediscovered in 2003. This small quarry was the only active one of the region in the end of the nineteenth
century. After clearing, the sedimentary succession of this quarry has been carefully prospected. In October
2004, an irregular stratum has been recognized at the bottom of the south side of the quarry. Among other
remains, mainly holoptychiid scales, this stratum has also provided a large lower jaw of the long-snouted
lungfish Soederberghia, a globally distributed Famennian genus known to be associated with tetrapods in
Greenland, Pennsylvania and Australia.
In spring 2005, the north side of the quarry was investigated and revealed a rich fossil-bearing
succession of differentiated strata. One of these layers provides the same unusual sedimentology as the sandstone
embedding the historical lower jaw, i.e., a flood facies composed of paleosoil clasts. This vertebrate-bearing
stratum, moreover showing the same rare sedimentology as the rock surrounding the tetrapod lower jaw,
suggests than the Ichthyostega-like tetrapod from Belgium comes from this stratum. An incomplete isolated
tetrapod maxilla was discovered in this very layer in 2007, confirming the presence of Devonian tetrapods at the
Strud locality.
Nevertheless, renewed interest in the Strud quarry requests the thorough studies of the identification of
a miospore assemblage for an accurate and certain dating, of the sedimentology and of the faunal and floral
composition of this tetrapod-bearing locality.
The succession of strata presents on the northern side of the quarry presents an almost vertical
inclination and is stratigraphically reversed. The different layers can be gathered in three main lithologies, from
bottom to top: a dolomitic sandstone, a mixed sandstone/siltstone and a fine-grained siltstone. This succession

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corresponds to a channel filling (Thorez, personal communication), i.e., accumulations of sediments and detritus
(plant, invertebrate, vertebrate fossil material) in a stream channel depending of the current dynamics. The top of
the section is composed of a greenish fine siltstone with complete and exceptionally well preserved crustacans
(Triops-like notostraceans, ostracods, and undetermined malacostraceans). This environment is considered as a
temporary pond, pointing the end of the flood event.
The composition of the sandstone surrounding the tetrapod lower jaw is very unusual (fluviatile
deposits generated by flood events). Such a lithofacies is usually known in the Evieux Formation in the northern
part of the Dinant synclinorium, Ardenne Allochthon, and the Namur Synclinorium, Brabant Parauthochthon
(Thorez et al. 1977; Bultynck & Dejonghe 2002) to which it was first attributed (Clément et al. 2004). The
palynological study is still in progress, but already it conclusively shows that the Strud locality is older than
previously thought (Clément et al. 2004; Blieck et al. 2007). According to Streel in Prestianni et al. (2007), it is
now considered to be upper middle Famennian in age (sensu Streel in Bultynck & Dejonghe 2002), i.e.
containing the lower part of the GF Miospore Zone and so close to but older than the East Greenland material.
This also means that the Strud quarry Evieux lithofacies might have the same age as the Souverain-Pré
Formation elsewhere (Bultynck & Dejonghe 2002, fig. 7). However, a more precise correlation between the
different Famennian sections of the Condroz area has still to be processed.
Although mainly disarticulated, fossil remains from Strud are regarded as autochthonous. The quality of
the material is surprisingly good, allowing accurate identifications, up to species level. Bone is well-preserved
and delicate elements such as Phyllolepis plates or dentary teeth on lower jaws are always found unbroken. The
exquisite preservation of arthropods and plants in the highest layers of the section, among which numerous seed
plant cupules representing the earliest seed plants worldwide (Prestianni et al. 2007), points to a reduced
postmortem transport, allowing to consider the taphonomic assemblage as a biological community.

2. Plant material

Fossil plants at Strud are known since Hock (1878) reported the presence of Rhacophyton in that small
quarry. In his comprehensive review of Belgian Famennian localities, Stockmans (1948) considerably expanded
the floral list.
Each lithology (i.e., dolomitic sandstone, a mixed sandstone/siltstone and a fine-grained siltstone)
contains plant remains, but not identical assemblages. All the specimens are compressions/adpressions, with no
anatomical details. Brief descriptions of the fossil plant contents for each lithology is given below:

The dolomitic sandstone: These deeply weathered beds contain many axes with apparent similar
morphology. They consist of large axes without fertile parts or vegetative endings (see Pl. 1, fig. 4 in Prestianni
et al. 2007). These show a very typical central ridge. They are attributed to the genus Rhacophyton, and belong
probably to the species Rhacophyton condrusorum. This genus represents one of the major components of the
Upper Devonian floras in Belgium.
The mixed sandstone/siltstone: These beds yield by far the largest diversity: zosterophylls, ferns,
progymnosperms and spermatophytes. Plant remains are however small and disarticulated. Zosterophylls are
represented by Barinophyton. From the stratigraphical point of view, this genus is one of the youngest
representatives of the group, as it went extinct during the early Carboniferous. One single specimen has been
found. It consists of an axis bearing two strobili. Ferns are rare; they are represented by Rhacophyton. In contrast
with a number of other Belgian localities (Fairon-Demaret 1996), Archaeopteris remains are rare at Strud (see
Pl. 1, fig. 1 in Prestianni 2007). The organisation of their axis and the shape of their leaves are characteristic of
the species Archaeopteris roemeriana (Göppert) sensu Stockmans, 1948. Two spermatophyte species have been
collected: Pseudosporogonites hallei and Moresnetia zalesskyi, which respectively correspond to the
Aglosperma and Moresnetia types of Prestianni (2005). Finally, many but dispersed pinnules of unknown
affinities have been found scattered in these beds (Pl. 1, fig. 2 and 5 in Prestianni et al. 2007).

The fine grained siltstone: This bed is very poor in fossil plants. Only two species have been found.
They were collected from two slightly different sediments. The coarser sediment contains well preserved cupules
of Moresnetia zalesskyi. These are here often found connected to the two or three distal most dichotomies of the
frond. The other sediment is fine-grained and better corresponds to a mudstone. It essentially contains Condrusia
rumex cupules. Two groups of sporangia have been collected from the same bedding plane. They are currently
under inverstigation. Their affinities are unknown, and those sporangia are left "incertae sedis". Nevertheless, on
the basis of their association with the seed plant Condrusia, they might represent the microsporangia of the latter.

3. Invertebrate material

The Strud locality seems to yield the oldest freshwater notostracean crustacans (Triops-like animal) as
well as undescribed and probably new taxa of malacostracean crustacans. All these exceptionally well preserved
specimens have been found, together with numerous ostracods, in the very fine siltstone-mudstone of the highest

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layers of the section. Some rare large isolated elements of eurypterid arthropods were also found in layers of the
mid-part of the section (i.e., the blackish mixed sandstone/siltstone rocks).
No marine invertebrate taxon has been found in this section so far.

4. Vertebrate material

Almost all the historical material of Famennian vertebrates from Belgium was found during the late
nineteenth century. Some specimens have been described and figured at this time (Lohest, 1888) and then fifty
years later (Leriche, 1931). More recently, some Coccosteid (Placodermi, Arthrodire) remains have been
described from the Famennian of Esneux (Lelièvre, 1982) as well as a new actinopterygian genus from the Early
Famennian of La Fagne (Taverne, 1997). The Paleozoic vertebrate fauna of northern France and Belgium has
been reviewed and published in 1995 (Cloutier and Candilier, 1995; Blieck and Lelièvre, 1995; Derycke et al.,
1995).
The Famennian vertebrate fauna of Strud has been reviewed based on historical and new material. A
brief overview of the composition of this Famennian Belgian fauna is reported here below:
Placoderms: The Antiarch Phyllolepis undulataLohest, 1888 is common in Strud. Isolated
anterolateral, median-dorsal and nuchal plates have been figured by Lohest (1888) and Leriche (1931).
Isolated median dorsal plates of the rare antiarch Groenlandaspis have been recently found in the
Paleontology collection of Louvain-la-Neuve, Belgium. This groenlandaspid material from Strud most probably
belongs to the same species than the recently described Groenlandaspis thorezi from Modave (Janvier and
Clément 2005). The genus Bothriolepis is very abundant in some Famennian quarries of Belgium (e.g.
Chèvremont, Liège Province and Spontin, Namur Province) but this antiarch seems to be rare in Strud (some
`Pterichthys' isolated plates housed in the IRScNB as well as anterior median dorsal plates and pectoral fin
elements recently found).
Acanthodians: Numerous isolated fin spines are present at Strud. These spines are slender, some of
them are smooth and others are ornamented with numerous parallel very fine ridges. No chondrichthyan or
gyracanthid remains has been found at Strud so far.
Actinopterygians: Some isolated remains of very small actinopterygians have been found in the
greenish fine siltstone/mudstone, some centimeters below the level where numerous complete crustaceans were
found. A preliminary study of these isolated scales, lower jaws, and cheek bones presumes that a single genus is
present at Strud.
Porolepiformes: Isolated Holoptychius scales and cleithra are very common in Strud. Complete
squamations have also been found although cranial bone plates or lower jaws are strikingly not so common.
Lungfishes: Isolated dipnoan dental plates are frequently found in the Late Devonian of Belgium. They
are all referred to the genus Dipterus but no detailed and accurate study of this material has been achieved.
Among the numerous different species of Dipterus established during the ninetheenth century, Leriche (1931)
has distinguished two different species, D. nelsoni Newberry, 1887 and D. fourmarieri Leriche, 1931. Both are
present in the Strud outcrop.
The long-snouted dipnoan Soederberghia is also present in Strud. An almost complete and large left
lower jaw is undistinguishable, in shape and size, of Soederberghia groenlandica from East Greenland (Clément
& Boisvert 2006). A quite abundant material of the smallest long-snouted lungfish Jarvikia is currently under
description. This latter genus was previously only known in East Greenland.
`Osteolepidids': Some rhombic, non-cosmine-covered, and tubercle-ornamented scales from Strud and
Modave (Destinez, 1904) have been referred to Glyptopomus (Glyptoloemus or Glyptolaemus) kinnairdi by
Lohest (1888). Glyptopomus is known in the Famennian of Scotland (Jarvik, 1950). Isolated small lower jaws
and scales, shiny cosmine-covered, could possibly belong to a megalichthyid osteolepiform.
Tristichopterids: Numerous isolated thin and rounded scales were found in Strud. They present a free
portion consisting of interconnected, sinuous ridges and their internal surface shows a drop-shaped knob. The
tristichopterid genus Eusthenodon is known to be present in the Famennian of Belgium as well as a larger
tristichopteridae gen. et sp. indet. (Clément, 2002; Clément et al. in press).
Rhizodontids: An isolated cleithrum from Strud was recently found in the Paleontology Collection of
Liège University. After preparation, the waisted and twisted shape of this small and complete left cleithrum
appears to be very characteristic of the rhizodontid group.
Among the rhizodontids, the shape of this cleithrum is very similar to that of the Late Devonian
Strepsodus? from Colombia (Janvier and Villarroel, 2000, pl. 5:8) and of Strepsodus sp. from Turkey (Janvier et
al., 1984, fig. 9A). This genus is known in the Carboniferous by Strepsodus sauroides and ?Strepsodus
anculonamensis from United Kingdom and from Queensland, Australia. This Famennian rhizodont cleithrum is
nevertheless the first occurrence of a Late Devonian rhizodontid from Europe.
Tetrapods: The Late Devonian tetrapod occurrences from Belgium consists nowadays only by an
incomplete right lower jaw and an incomplete left maxilla. The lower jaw, found in Strud during the nineteenth

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century and housed in the historical collection of the Liège University, has been referred to an Ichthyostega-like
form (Clément et al., 2004). The genus Ichthyostega was previously only known in the Late Famennian of
Greenland (Jarvik, 1996; Marshall et al., 1999; Blom 2005). The fragment of maxilla was found in 2007 in the
sandstone layer presenting the same unusual sedimentology than that of the historical lower jaw.

5. Environment

The different lithologies provide environmental as well as climatic information. Globally, the channel
filling matches to an arid climate interrupted by occasional heavy rainfalls of short duration.
Flooding rains probably reworked the sediment and, at the same time, the surrounding vegetation, in
this case a monotypic Rhacophyton population. This suggests a general dry climate with regular abundant
rainfalls. The conglomeratic siltstone and its fossiliferous content have been deposited in two phases. Firstly, low
energy sedimentation allowed the deposition of fine grained, fossil rich silt. Later, flooding rains reworked this
silt in more sandy sediment. Such depositional conditions allowed the mixing of plant fossils and explain the
observed diversity as they probably fossilized in different environments as well as at different moments of the
year. The finer grained sediment corresponds to the last stage of the channel filling. It was accumulated in a very
low energy river or in a calm but abandoned channel. Lycopsids are notably absent from the collected material.
By contrast with East American localities, this is a general characteristic for the Upper Devonian of Belgium
(Fairon-Demaret 1996). Streel and Marshall (2006) have suggested that the Late Devonian emerging Variscan
Mountains blocked the humidity of trade winds and induced aridity in Belgium. As most lycopsids need water to
complete their life cycle, it is suggested that aridity might have disadvantaged them. This might also explain why
a single species of Archaeopteris is known in Belgium (A. halliana/roemeriana), although the genus was
extremely common. Five species are for example known to occur in USA.
The Upper Devonian mega-environment in Belgium was a large lagoonal system developed along the
south of the Euramerican continent. Aridity led to the development of large evaporitic areas. The information
collected at Strud conforms to those conclusions at a smaller scale and demonstrate the Upper Famennian
environments were diversified and included a large range of ecosystems. This favored the plant diversity
partially recorded in the second bed of Strud.

6. Vertebrate paleogeography

Devonian tetrapods seem to present a high level of endemism although the Ichthyostega-like form from
Strud suggests that they were not as restricted as previously thought (Clément et al. 2004).
Lohest (1888, pl. VIII: 2, 5) misinterpreted a Devonian tetrapod lower jaw as a large fish remain and
assigned it to a new species of Dendrodus, D. Traquairi. This fragment of mandible shows typical tetrapod
characters and looks very similar to the corresponding jaw portion of Ichthyostega from Greenland6. Although
the find cannot be referred with certainty to Ichthyostega, it is strong evidence for the existence of a close
relative of this tetrapod genus outside Greenland.The biogeographical distribution of Devonian tetrapods is near-
global but extremely restricted (to a single locality or small geographic area) at the generic level, which was
presumably a result of the colonization of continental environments that were less favourable to wide dispersal
than were marine ones (Blieck et al. 2007). The occurrence of an Ichthyostega-like form in Belgium extends the
geographical distribution for at least one taxon and suggests that they were not as restricted as previously
thought.
The vertebrate assemblage of the Strud site is important for the understanding of paleobiogeographic
relationships of Laurussia with Gondwana. The lungfish Soederberghia is only known in the Late Frasnian and
Lower Famennian of Australia, in the Late Famennian of East Greenland and Pennsylvania (Ahlberg et al.
2001), and now in the Late Middle Famennian of Belgium. The Gondwanan native Phyllolepis is typical of the
Famennian stage in Laurussia (East Greenland, Pennsylvania, Russia, and Belgium) (Young 2003). The first
Euramerican taxa of the Rhizodontida, a clade of tetrapodomorph fishes native from Eastern Gondwana
(Johanson 2004), are only known in the Famennian tetrapod-bearing localities of Pennsylvania (Davis et al.
2001, 2004) and now Belgium.
The first dispersal of the Phyllolepida from South China to North-Eastern Gondwana has been dated at
the Pragian--Emsian boundary, during the E'Em bioevent of Walliser (1995) (Dupret & Zhu 2008). Indeed,
before this bioevent, the endemism rate of Chinese early vertebrates is very high; then, because of the invasion
of and the subsequent competition with Gondwanan taxa, this endemism rate falls drastically (e.g. Galeaspida).
The second major dispersal episode of the Phyllolepida occurs along the northern margin of the
Gondwana, westward until Middle East (Turkey) and northern South America (Venezuela). The recent
Phyllolepis occurrence from the Middle-Late Devonian of Venezuela (Young et al. 2000), as well as the
rhizodontid occurrences in the Famennian of Colombia and Turkey (Janvier and Villarroel 2000; Janvier et al.

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1984), strengthens the hypothesis of a nonmarine dispersal episode across the northern Gondwana margin during
the Middle/Late Devonian (Young 1990).
The third dispersal episode is dated at the level of the Frasnian--Famennian boundary, between
Gondwana and Euramerica. By that time, the Placodermi Phyllolepididae and Groenlandaspididae, and the
Sarcopterygii Rhizodontidae and Megalichthyidae and the Dipnoi Soederberghia would have migrated
northward to Euramerica, while the Antiarch Asterolepididae and the Sarcopterygii Holoptychius migrated
southward to Gondwana. The most probable area of "transfert" between the two landmasses is the North of
South America (Clément et al. 2005; Dupret et al. 2005).
All the other taxa found in Strud present a high level of cosmopolitanism (e.g., Bothriolepis,
Holoptychius, `Dipteridae', Glyptopomus, large derived tristichopteridae) and are found in the Late Devonian of
Laurussia and Gondwana but not in China. For example, Bothriolepis and Holoptychius, respectively the most
widespread placoderm and sarcopterygian, are known in almost every Upper Devonian vertebrate-bearing
localities of the world.
The relative position of the two supercontinents Gondwana and Laurussia during the Late Devonian
was first supposed to be widely separated from each other on the basis of paleomagnetism evidence (e.g., Van
der Voo 1988; Kent and Van der Voo 1990; Li et al. 1993). Another hypothesis, based on paleomagnetism but
also on biogeography and facies distributions, supposed that the two supercontinents were in close contact
during the Devonian times (e.g., Scotese and McKerrow 1990; McKerrow et al. 2000; Ahlberg et al. 2001;
Johanson, 2004; Young 1990, 2003).
The vertebrate assemblage of Strud, and relationships of the various Famennian vertebrate fauna of the
world, contradict the previous hypothesis of a wide separation between Laurussia and Gondwana. On the
contrary, they reinforce the hypothesis of a continental connection at or near the Frasnian-Famennian boundary
between the southeastern margin of Laurussia and northern margin of Gondwana.

7. References

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Geologica Belgica, v. 8, pt. 1-2 (in press).
JANVIER, P., & VILLARROEL, C. 2000. Devonian vertebrates of Colombia: Palaeontology, v. 43, pt. 4, p. 729-763.
JANVIER, P., LETHIERS, F., MONOD, O., & BALKAS, Ö. 1984. Discovery of a vertebrate fauna at the Devonian-Carboniferous
boundary in SE Turkey (Hakkari Province): Journal of Petroleum Geology, v. 7, pt. 2, p. 147-168.

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JARVIK, E. 1950. On some osteolepiform crossopterygians from the Upper Old Red Sandstone of Scotland: Kunglinga Svenska
Vetenskapsakademiens Handlingar, ser. 4, v. 2, pt. 2, p. 1-35.
JARVIK, E. 1996. The Devonian tetrapod Ichthyostega: Fossils and Strata, v. 40, p. 1-213.
JOHANSON, Z. 2004. Late Devonian sarcopterygian fishes from eastern Gondwana (Australia and Antarctica) and their importance in
phylogeny and biogeography, in Arratia, G., Wilson, M.V.H., andCloutier, R, eds., Recent advances in the origin and early
radiation of Vertebrates: München, Verlag Dr. Friedrich Pfeil, p. 287-308.
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12, p. 49-56.
LELIEVRE, H. 1982. Ardennosteus ubaghsi n.g., n. sp. Brachythoraci primitif (Vertébré, Placoderme) du Famennien d'Esneux (Belgique) :
Annales de la Société Géologique de Belgique, v. 105, p. 1-7.
LERICHE, M. 1931. Les Poissons Famenniens de la Belgique: Mémoires de la Classe des Sciences de l'Académie Royale de Belgique, v.
10, pt. 5, p. 1-72.
LI, Z.X., POWELL, C.M., & TRENCH, A. 1993. Palaeozoic global reconstructions, in Long, J.A., ed., Palaeozoic Vertebrate
Biostratigraphy and Biogeography: London, Belhaven, p. 25-53.
LOHEST, M. 1888. Recherches sur les poissons des terrains paléozoïques de Belgique. Poissons des Psammites du Condroz, Famennien
supérieur: Annales de la Société Géologique de Belgique, mém. 15, p. 112-203.
MARSHALL, J.E.A., ASTIN, T.R., & CLACK, J.A. 1999. East Greenland tetrapods are Devonian in age: Geology, v. 27, no. 7, p. 637-640.
MCKERROW, W.S., MAC NIOCAILL, C., AHLBERG, P.E., CLAYTON, G., CLEAL, C.J., & EAGAR, R.M.C. 2000. The Late
Palaeozoic relations between Gondwana and Laurussia, in Franke, W., Haak, V., Oncken, O., and Tanner, D., eds., Orogenic
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PRESTIANNI, C. 2005. Early diversification of seeds and seed-like structures. In: Steemans, P., and Javaux, E. (eds.) Pre-Cambrian to
Palaeozoïc Palaeopalynology and Palaeobotany. Carnets de Géologie / Notebooks on Geology, Memoir 2005/02, Abstract 06.
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2007/01, Abstract 07.
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Bruxelles, vol. 110, 84 p.
STREEL, M., & MARSHALL, J. 2006. DevonianCarboniferous boundary global correlations and their paleogeographic implications for
the Assembly of Pangaea. In: The XVth International Congress on Carboniferous and Permian Stratigraphy, 481-496.
TAVERNE, L. 1997. Osorioichthys marginis, "Paléonisciforme" du Famennien de Belgique, et la phylogénie des Actinoptérygiens
dévoniens (Pisces) : Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre, v. 67, p. 57-78.
THOREZ, J., DREESEN, R., & STREEL, M. 2006. Famennian. In: Dejonghe L. (ed.), Current status of chronostratigraphic units named
from Belgium and adjacent areas. Geologica Belgica, Bruxelles, vol. 9, n° 1-2, p. 27-45.
THOREZ, J., STREEL, M., BOUCKAERT, J. & BLESS, M. J. M. 1977. Stratigraphie et paléogéographie de la partie orientale du
synclinorium de Dinant (Belgique) au Famennien supérieur : un modèle de bassin sédimentaire reconstitué par analyse
pluridisciplinaire sédimentologique et micropaléontologique. Mededelingen van de Rijks Geologische Dienst, N.S., 28 (2), 17-28.
VAN DER VOO, R. 1988. Paleozoic paleogeography of North America, Gondwana, and displaced terranes: comparisons of paleomagnetism
with paleoclimatology and biogeographical patterns: Geological Society, bull. 100, p. 311-324.
WALLISER, O. H. 1995. Global events in the Devonian and Carboniferous; pp. 225-250 in O. H. Walliser (ed.), Global events and event
stratigraphy in the phanerozoic. Springer, Berlin, Heidelberg & New York.
YOUNG, G. C. 1981. Biogeography of Devonian vertebrates. Alcheringa, 5, 225-243.
YOUNG, G.C. 1990. Devonian vertebrate distribution patterns and cladistic analysis of palaeogeographic hypotheses, in McKerrow, W.S.,
and Scotese, C.F., eds., Palaeozoic Palaeogeography and Biogeography: Geological Society, London, Memoir no. 12, p. 243-255.
YOUNG, G.C. 2003. North Gondwanan mid-Palaeozoic connections with Euramerica and Asia: Devonian vertebrate evidence: Courier
Forschunginstitut Senckenberg, no: 242, p. 169-185.
YOUNG, G.C., MOODY, J.M., & CASAS, J.E. 2000. New discoveries of Devonian Vertebrates from South America, and implications for
Gondwana-Euramerica contact: Compte-rendus de l'Académie des Sciences, Paris, v. 331, p. 755-761.

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Appendix. List of the biota of the Famennian locality of Strud, Belgium

FLORA

Filicopsids:

Rhacophyton sp.
Progymnosperms:
Archeopteris roemeriana
Gymnospermes:
Condrusia rumex

Pseudosporogonites hallei

Moresnetia zalesskyi
Zosterophyllopsids:
Barinophyton condrusorum
Incertae sedis:
Cf. Calathiops sp.
Sphenopteris flaccida
Sphenopteris modavensis

INVERTEBRATE FAUNA

Crustacea

Notostraca
Triops-like tadpole shrimp

Ostracoda
smooth ostracods indet.

Malacostraca
large specimens indet.

Eurypterida

large Eurypterida indet.

VERTEBRATE FAUNA
Placodermi

Arthrodira
Phyllolepis undulata
Groenlandaspis thorezi

Antiarcha

Bothriolepis sp.
Acanthodii

acanthodian indet.
Actinopterygii

Actinopterygian indet.
Sarcopterygii
Dipnomorpha
Porolepiformes

Holoptychius flemingi

Holoptychius nobilissimus
Dipnoi

`Dipterus' sp.

Rhynchodipteridae

Soederberghia cf. groenlandica

Jarvikia sp.

Tetrapodomorpha

'Osteolepiformes'

Glyptopomus sp.

Cosmine-covered osteolepidid (?Megalichthyidae)
Tristichopteridae indet.

Rhizodontida

Rhizodontida indet.

Tetrapoda

Ichthyostega-like form

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Plate I

Figure 1. Location of the Strud locality in Belgium.
Figure 2. The Belgian paleontologist
and paleoanthropologist Maximin
Lohest, from the Liège University,
described numerous Paleozoic
vertebrates from Belgium as well as
prehistoric remains (Neanderthalians
from the Spy cave).

Figure 3. Precise location of the Strud village in the Namur Province.

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Plate II

Figure 1. Archaeopteris roemeriana (GÖPPERT) STOCKMANS, ULg n° 15201, scale: 0.5 cm.
Figure 2. Sphenopteris modavensis STOCKMANS 1948, ULg n° 15221, scale: 1 mm.
Figure 3. Condrusia rumex STOCKMANS, ULg n° 15222, scale 1 mm.
Figure 4. Rhacophyton sp. CRÉPIN, ULg n° 15203, scale 1 cm.
Figure 5. Sphenopteris flaccida STOCKMANS 1948, ULg n° 15215, scale 5 mm.
Figures 6-7. Moresnetia zalesskyi STOCKMANS, ULg n° 15216 and 15217, scales 0.5 mm.
(from Prestianni et al. 2007)

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Plate III

Figure 1. Placoderm Phyllolepis undulata (from Leriche 1931), x1.
Figure 2. Placoderm Bothriolepis sp., scale: 1 cm.
Figure 3. Left lower jaw of the long-snouted lungfish Soederberghia (from Clément & Boisvert 2006).
Figure 4. Left lower jaw of a small actinopterygian, scale 1mm.
Figure 5. Right cleithrum of a small tetrapodomorph rhizodontid.
Figure 6. Right lower jaw of an Ichthyostega-like tetrapod (from Lohest 1888; Clément et al. 2004), scale 1 cm.
Figure 7. Isolated scale of the large porolepiform Holoptychius.
Figure 8. Complete Triops-like notostracan crustacean, scale: 1 mm.

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Plate IV

Figure 1. Lithostratigraphical subdivsions of the Famennian Stage within the Dinant Synclinorium. Dashed lines
correspond to successive oolitic ironstone levels (event-stratigraphical marker beds) (from Thorez et al. 2006).

Figure 2. Paleogeographical maps of the Devonian and Late Carboniferous times. Stars correpond to the
position of Belgium (from the website "Une introduction à la géologie de la Wallonie":
http://www2.ulg.ac.be/geolsed/geolwal/geolwal.htm).

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TAPHONOMY AND SEDIMENTOLOGY OF THE `BLACK MARBLE'
OF DENÉE, A FOSSIL CONSERVATION DEPOSIT FROM THE VISÉAN
(MISSISSIPPIAN) OF BELGIUM

Bernard MOTTEQUIN

Unité de Paléontologie animale et humaine, Université de Liège, Bât. B18, Allée du 6 août, B 4000 Liège 1,
Belgium, bmottequin@ulg.ac.be

1. Introduction

The localities of Tournai and Visé, respectively the historical type areas of the Tournaisian and Viséan
stages (Hance et al. 2006a-b), have contributed to the fame of the Lower Carboniferous of the Namur-Dinant
Basin [southern Belgium and northern France (Avesnois)] by the great diversity and the abundance of the
macrofaunas (see references in Demanet 1958). Nevertheless, their study (except rugose corals and trilobites)
has been neglected for many years. In addition to both these well-known fossiliferous localities, the quarries
located around the village of Denée (Namur province) (Figure 1) have yielded remarkably preserved but rare
fossils (including echinoderms and fishes), which have been collected within the `black marble' of Denée, a
black coloured limestone of early Viséan age.

Figure 1. General context of Lower Carboniferous sedimentation in north-western Europe showing the
distribution of emergent areas and Waulsortian mounds at the end of the Tournaisian [modified from Ziegler
(1990) and Devuyst & Dehantschutter (2007)]. B = Basin; D = Denée.

All the fossils, with few exceptions, were collected at the end of the 19th century and at the beginning of
the 20th century by quarrymen when the `black marble' was intensively and manually quarried. Most of the
quarries were subterranean and, nowadays, most of them are disused and flooded. If it had not been worked, the
`black marble' of Denée would have been considered probably as azoic due to the rarity of the fossils. The bulk
of the material is deposited at the Maredsous abbey (`Centre Grégoire Fournier'), but additional specimens are
housed in the University of Liège, the Royal Belgian Institut of Natural Sciences (Brussels), and the Museum of
Comparative Zoology (Harvard). A modern systematic revision is urgently needed for most of the invertebrate
phyla; the latest comprehensive list of the faunas dates back to Fournier and Kaisin (1929). As is generally the
case with old collections, the origin of the specimens is usually not known with precision, except for some
fossils which have a mention of the quarried level (`la Veine', `les Drîs') on their label. We can suspect that most

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of the specimens were collected at Denée, from the exploited levels figured by Fournier (in Fournier & Pruvost
1928) (Figure 2).

Figure 2. Distribution of fishes within the `black marble' of Denée (Molignée Formation) as exposed in the
quarries of the Denée area [modified from Fournier in Fournier & Pruvost (1928)].

Figure 3. Late Tournaisian sedimentation areas in the Namur-Dinant Basin (not palinspastic) (modified from
Poty et al. 2006). 1. Denée; 2. Salet; 3. Sovet. S.A. = sedimentation area.

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Figure 4. Evolution of the Condroz sedimentation area (CSA) and the Dinant sedimentation area (DSA) during
the third-order sequences 4 (A), 5 (B) and 6 (C) (numbered black arrows) [Tournaisian (Ivorian) to Lower
Viséan (Moliniacian)] (modified from Hance et al. 2001); see also Figure 3. The Banc d'Or de Bachant is a
bentonite, locally transformed into a palaeosol (Delcambre 1989). The Longpré Formation includes, from base to
top, the Flémalle and the Avins members; the Braibant Member corresponds to the top of the Sovet Formation
(Poty et al. 2002). Mbr: Member; MOL*: Molignée Formation (`black marble' of Denée); MOL**: Molignée
Formation (`black marble' of Dinant).

2. Geological setting

The Namur-Dinant Basin developed on the SSE margin of the London-Brabant Massif (Figures 1 and 3)
in a back-arc extensional setting (Hance et al. 2001), north of the Ligerian Arc (e.g. Leeder 1988). In the course
of late Tournaisian and early Viséan time, the ramp setting that has prevailed since the early Tournaisian
progressively evolved to a rimmed-shelf and to a broad flat-topped platform of regional extent during the middle
and late Viséan (Hance et al. 2006b). On the basis of their lithostratigraphic character, several sedimentation
areas have been defined within the Namur-Dinant Basin by Poty (1997) and Hance et al. (2001). These are
(Figures 3 and 4): 1) the Hainaut sedimentation area was an area in which subsidence allowed accumulation of
about 2500 m of Lower Carboniferous rocks, including several thick evaporitic intercalations; 2) the Namur
sedimentation area (NSA) characterized by the more proximal facies and the less complete lithostratigraphic
succession; 3) the Condroz sedimentation area (CSA) with relatively proximal facies and some sedimentary
breaks it displays southward and southwestwards a transition with the Dinant sedimentation area; 4) the Dinant
sedimentation area (DSA) displaying the deepest water facies with the development of carbonate mounds

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characterised by microbially mediated muds (= Waulsortian mounds; see references in Lees 2006), as well as an
almost complete stratigraphic succession; 5) the southern Avesnois sedimentation area (ASA) showing a similar
situation to that of the CSA, but with a markedly different lithostratigraphy (shallower water facies); 6) the Visé-
Maastricht sedimentation area suffered block faulting during the Devonian and Mississippian. It was connected
with the NSA during the Upper Devonian and Tournaisian and evolved to a graben that was open to the Campine
Basin during the Viséan (Poty 1997). In Belgium, the absence of Lower Carboniferous exposures south of the
DSA, due to post-Variscan erosion, does not allow the precise delineation of the eastern extension of the ASA.
However, according to recent investigations of Pirotte (2006), it seems, at least, that the ASA borders the
southern margin of the DSA, south of the Waulsortian complex (Figure 5).

Figure 5. Pirotte's (2006) modified model of stacking of the third-order sequences (numbered black arrows) in
the southern part of the Dinant sedimentation area and in the southern Avesnois sedimentation area, from the
Lower Tournaisian to Lower Viséan; see also Figure 1. DEN = `black marble' of Denée; DIN = `black marble'
of Dinant; Lef. = Leffe.

The `black marble' of Denée was formerly included in the `black marble' of Dinant (`V1a' of the
Belgian authors), but according to Conil (1967), it precedes this latter unit, strictly speaking, on the basis of the
foraminifer associations (Figures 4 and 5). Now, both former units are included in the diachronic Molignée
Formation (sensu Poty et al. 2002) of early Viséan age [regional Moliniacian Substage (e.g. Devuyst et al.
2006)]. This formation is developed only in the central part of the DSA between the prograding platform and the
Waulsortian complex (Hance et al. 2001) running along the border between the DSA and the ASA (Figures 35).
The Molignée Formation consists of a succession of thin-bedded (less than one metre to several metres thick),
commonly laminated black limestones which alternate with thick-bedded, dark-grey limestones (`thick beds')
(Figure 6). These alternations correspond to the `polysequences' and `monosequences' of Mamet (1964). The
contacts between both lithotypes are always clear-cut.
The range of the `black marble' of Denée, in terms of Mississippian Foraminifer Zone (MFZ), spans the
interval of the MFZ 9 (upper part) to the MFZ10 according to Devuyst and Hance (in Poty et al. 2006). In the
Salet road section (stratotype of the Molignée Formation), the `black marble' of Denée begins at bed 191 and
ends at bed 273 (Figure 6A); 39 m thick (see Hance (1988), Devuyst et al. (2006) and Poty et al. (2006) for
detailed logs of this section). Thus, it does not correspond completely to the Molignée Formation (from bed 169
to the top of bed 294; c. 58 m thick) as redefined by Poty et al. (2002).

3. The `black marble' of Denée, a fossil conservation deposit

Besides numerous ichnofossils (crawling and grazing traces), the `black marble' of Denée has yielded
the following biota: chondrostean and elasmobranch fishes (e.g. Traquair in de Koninck 1878; Pruvost in
Fournier & Pruvost 1922, 1928; Woodward 1924; Derycke et al. 1995, Maisey 2007) (Figures 2, 7AB),
echinoids (e.g. Jackson 1929) (Figure 7CD), ophiuroids (Fraipont 1904) (Figure 7E), crinoids (Fraipont 1904)
(Figure 7F), holothuroids (?), dendroid graptolites (e.g. Ubaghs 1941) (Figure 7GH), rugose (Figure 7I) and

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tabulate corals, sponges, conulariids, bryozoans, brachiopods (Delépine 1928) (Figure 7J), phyllocarids,
trilobites, bivalves (Demanet 1929) (Figure 7K), gastropods, goniatites (Delépine 1940), nautiloids, and floated
plant remains. Most of the fossils belong to the autochthonous epibenthos (Mottequin 2004). The echinoid fauna,
including the largest and most remarkable Palaeozoic specimens, as well as the fishes, have contributed largely
to its fame. Nowadays, it is not possible to sample all the fauna anymore because of the poor exposures of the
Molignée Formation. Palaeoecology and taphonomy were studied by Mottequin (2004).

Figure 6. A. Partial log of the Salet road section (stratotype of the Molignée and Salet formations) (modified
from Poty et al. 2006). Bio. = Biostratigraphy; Chrono. = chronostratigraphy; Litho. = lithostratigraphy; MFZ =
Mississippian Foraminifer Zones of Devuyst & Hance (in Poty et al. 2006). B. Molignée Formation in its
stratotype showing the alternating thicker bedded and thinner bedded units. Bed numbers are those of Overlau
(1966). B = `black marble' facies; T = `thick beds' facies.

During the Moliniacian, the colonization of the sea floor of the central part of the DSA by the benthos
and the diversity of the latter were strongly influenced by oxygen concentration. Only some organisms were able
to develop in this particular environment. Most of the bivalves belong to the `paper pecten' morphotype (Figure
7K) which is diagnostic of dysaerobic environments according to Allison et al. (1995). Echinoderms, especially
the echinoids, seem to have been well adapted to face this poorly oxygenated environment, as indicated by their
relative abundance in the collection and their preservation in situ (complete tests with spines in anatomical
connection). This is not surprising because these organisms are known in upper dysaerobic facies of modern
oxygen deficient basins; Savrda et al. (1984) reported also ophiuroids, holothuroids, polychaetes, crabs and
gastropods in their lower dysaerobic facies. Among brachiopods, representatives of the suborder Productidina

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predominate in the collection. Some of them have spines of much greater length than the shell bearing them
(Delépine 1928) (Figure 7J). These long spines were most probably for support (Brunton & Mundy 1988) and
acted as a stabilizing snowshoe to prevent shells from sinking in the muddy substrate (Bowen et al. 1974). Their
preservation attests to the absence of or only minor transport. Productidina would indicate better oxygenated
conditions than those prevailing during the colonization by bivalves and echinoids.

Figure 7. Fossils of the early Viséan-aged `black marble' of Denée (Molignée Formation). A. Benedenius
deneensis Traquair; B. Denaea fournieri Pruvost; C. Proterocidaris giganteus de Koninck; D. Fournierechinus
deneensis Jackson; E. Taeniaster? fournieri Fraipont; F. Graphiocrinus longicaudatus (Fraipont); G.
Ptiograptus fournieri Ubaghs; H. Dictyonema fraiponti Ubaghs; I. Caninophyllum sp.; J. unidentified productid
brachiopods; K. `Pterinopecten' dumontianus (de Koninck). Scale bars are 5 cm for AD and 1 cm for EK.

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The `black marble' of Denée is a fossil conservation deposit (`fossil Lagerstätte'; see discussion in
Shields 1998), i.e. the skeletons and the tests of organisms are preserved in their entirety. However, soft parts
have not really been highlighted with certainty although Van Straelen (1926) described an enigmatic fossil under
the name of Medusina boulengeri that he interpreted as the mould of the exombrella of a jellyfish. It has been
assigned to the Medusae incertae sedis by Harrington & Moore (1956), but Mottequin (2004) reinterpreted it as
a probable burrow. This hypothesis must be confirmed by X-rays analysis. Exceptional burrows have been
recognized within the `black marble' facies such as Zoophycos. Rapid burial is considered here as one of the
major factor of preservation [`obrution deposits' of Seilacher et al. (1985)]. It is well exemplified by the
excellent preservation of most of the echinoderms. Low oxygen concentrations prevailing within the substrate
slowed down the disarticulation of the tests and excluded the eventual predators.
Carpentier (1913) and Jackson (1929) have compared the `black marble' of Dinant with the lithographic
limestone of Portlandian age from the Solnhofen area in Germany which is the type of the `taphofacies IF:
dysoxic/anoxic basin' of Brett et al. (1997) based on echinoderm taphonomy. However, in the case of Denée, the
organisms are essentially benthic and generally autochthonous, contrary to the lagoon of Solnhofen, where the
benthos was imported during storms and only survived for some days before the re-establishment of hypersaline
conditions. The `black marble' is close also to the to obrution deposit typified by the Hunsrück Slate of Emsian
age (Brett & Seilacher 1991) and more especially by the locality of Budenbach. Although the faunas of the
`black marble' of Denée are not pyritized, the burial of the organisms in the Belgian conservation fossil deposit
is approximately the same as in the German locality (Sutcliffe et al. 1999). Nevertheless, the environment in
Denée may have been less oxygenated as indicated by the presence of bivalves of the `paper pecten' morphotype
and the very rare to absent trilobites.

4. Sedimentation in the central part of the part of the Dinant sedimentation during the early Moliniacian

4.1. General context

An important factor that influenced the sedimentation and the palaeoenvironment of the Molignée
Formation was the topographic context inherited from the upper part of the Tournaisian. During the Ivorian
(Tournaisian), the DSA was characterized by the build-up of large Waulsortian mounds whose maximal
development gave rise to a discontinuous barrier in its south-western part (Waulsortian complex) (Lees 1997).
At the same time, the ASA recorded a subsidence rate lower than the one of the DSA, from which it was
separated by a synsedimentary fault (Pirotte 2006). The end of the Tournaisian, i.e. at the end of the third-order
sequence 4 of Hance et al. (2001), is marked by a major sea-level drop which induced the emersion of the
shallower areas of the Namur-Dinant Basin (karstification of the top of the Avins Member) and the southward
progradation of the shelf during the next sequence (sequence 5) (Figure 4B). In the early Viséan (sequence 5)
that corresponds to the period of deposition of the Molignée Formation, the central part of the DSA evolved as a
residual intra-platform basin bounded by the prograding shelf to the north and the Waulsortian complex built
against a major synsedimentary fault to the south. The eustatic magnitude of this sequence was probably low,
because, in the Namur-Dinant Basin, it is only recorded in the DSA. In the ASA, the Terwagne Formation
(sequence 6) lies directly on the Godin Formation (sequence 4) whereas in the CSA and the NSA, the former
caps the Avins Member of the Longpré Formation (Hance et al. 2001) that indicates clearly a sedimentary gap
(Figure 3). According to Devuyst (2006), it is probable that the top of the Waulsortian complex emerged also,
because it was growing in very shallow water at the end of the Tournaisian (Lees 1997), although no trace of
such emersion has, however, yet been found. According to Devuyst (2006), this may be the result of the
monotonous facies which hinder the recognition of facies variations.

4.2. Microfacies of the Molignée Formation

Among the thin beds (`black marble' facies), the dominant microfacies consists of well-sorted and
laminar packstones to grainstones composed of calcispherids, peloids, moravamminids, plurilocular
foraminifers, algal fragments and other allochems of similar size (Figure 8BC). By their characteristics (flat
parallel laminations, allochems of small dimensions), they have been interpreted previously as distal turbidites
by Overlau (1966), which poured out on the marine floor and were intercalated with mudstones with
calcispherids and radiolarians (Figure 8A). The packstones-grainstones shift laterally to wackestones-packstones
due to the progressive decrease of the turbidity currents. Moreover, the rugose and tabulate corals recovered in
the `black marble' are always as reworked fragments which are transported into the basin via the turbidity
currents. Numerous skip marks produced by broken shells of gastropods and cephalopods also confirm the
existence of bottom currents. The burrows are exceptional (Zoophycos sp.; see also section 3). The `thick beds'
are mainly composed of bioturbated wackestones to packstones with a high diversity of allochems, among which
include as major components: bryozoans, echinoderms, gastropods, ostracods, trilobites, and foraminifers

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(especially Tetrataxis) (Figure 8E). Sponge spicules, algae, Rectangulina, Globochaete, cephalopods, solitary
rugose corals (Cyathaxonia sp.), brachiopods, and calcispherids have been also recognized (see section 4.3 for
their interpretation). Rudstones (Figure 8D) and grainstones occur at the base of some beds. The microfacies of
the `thick beds' occurring in the Molignée Formation are reminiscent of those of the underlying Leffe Formation
(Noël in Groessens & Noël 1975), though coated clasts and oncoids are less abundant. According to Lees et al.
(1977), the carbonate mud of the Leffe Formation is probably derived from the Waulsortian mounds developed
in the DSA. However, the Leffe facies has never been identified with certainty in other areas with Waulsortian
mounds (Lees 1997). Devuyst (2006) suggested that a large part of the lime mud of the upper part of the Leffe
Formation was exported from the mounds when they reached depths at which the accommodation space for
aggradation was reduced.

Figure 8. Microfacies of the Molignée Formation in the Denée area; the bed numbers are those of Mottequin
(2004). AC, `black marble' facies. A. Alternation of mudstone, wackestone and packstone with radiolarians, En
Gilotia locality in Maredret (bed 8a). B. Well-sorted, laminated grainstone with peloids and calcispherids,
Debras quarry in Salet (bed 49). C. Grainstone with peloids and calcispherids, and packstone with bioclasts,
Debras quarry in Salet (bed 2e). DE, `thick beds' facies. D. Rudstone with lithoclasts, crinoids, bryozoans and
peloids, Debras quarry in Salet (bed 27a). E. Wackestone with peloids, trilobites, crinoids, bryozoans and mud
balls, En Gilotia locality in Maredret (bed 24). All scale bars are 10 mm.

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4.3. Palaeoenvironmental interpretation of the Molignée Formation in the Denée-Salet area

The origin of the alternations of `black marble' and `thick beds' facies has been previously interpreted
by Mottequin (2004) as shallowing-upwards parasequences with the `thick beds' at their base and the `black
marble' facies at their top (Figure 6B); their origin would have been linked to the periodic confinement of the
DSA due to sea-level fluctuations. Low oxygen concentrations are suggested by the remarkable preservation of
the fauna and the existence of dysaerobic organisms such as the bivalves of the `paper pecten' morphotype.
Because of the required depth conditions for the expansion of anoxia in Recent confined basins (Murray et al.
1989), Mottequin (2004) suggested that the `black marble' facies would have been developed between 100 and
150 m water depth, whereas the `thick beds' would have been deposited between 150 and 200 m depth. This last
estimation used the presence of the plurilocular foraminifer Tetrataxis within the `thick beds' which would
indicate a depth less than 200 m, according to Lees (1997), as well as the reconstruction of the Namur-Dinant
Basin by Hance et al. (2001).
The confinement of the central part of the DSA was probably due to minimal sea-level fluctuations,
partly induced by glacio-eustasy (see discussion in Mottequin 2008). `Spasmodic' synsedimentary tectonics of
the basin may have also influenced the palaeoenvironment. Indeed, two major synsedimentary faults have been
inferred previously on the basis of thickness variations of some formations, as well as on sudden facies changes.
The first one would have been located in the Sovet area (Figure 4B), but it is believed to have been active at least
after the sedimentation of the Braibant Member forming the cap beds of the Sovet Formation (Hance 1988;
Hance et al. 2001; Devuyst 2006; Poty et al. 2006). The second fault delimited the southern part of the DSA and
the northern part of the ASA (Pirotte 2006; Figure 5); it appeared probably at the end of the Hastarian when the
ASA began to differentiate from the DSA with the appearance of shallower water facies in the former. If both
were active during the deposition of the Molignée Formation, the DSA would have thus evolved as a subsiding
zone.
Devuyst (2006) recognized three large-scale shallowing-upwards cycles in the stratotype of the Sovet
Formation (the lateral equivalent of the Molignée Formation) which he interpreted as reflecting the southwards
progradation of the platform margin as successive mega-clinoforms. So, most of the turbidites occurring within
the `black marble' facies would originate from the northern part of the DSA, as previously suggested by Hance
et al. (2001), on the basis of the allochems present in the packstones-grainstones. The turbidites would result
from violent storms, earthquakes or clinoform instability due to an oversteepened slope.
The persistence within the Molignée Formation of facies similar to those observed in the Leffe
Formation can be partly explained by the fact that the abrasion of the Waulsortian mounds continued after their
decay they stopped growing in the late Tournaisian and also by the fact that the topography inherited from
these buildups remained for a while. Waulsortian mounds are recognized in the Denée-Salet area (e.g.
Delcambre & Pingot 2004).
Because of the random mode of dispersion of the turbidites, it is very difficult to correlate the various
sections exposing the Molignée Formation in the Denée-Salet area. The microfacies recognized in Denée and
Salet are different because Salet occupied a more proximal position. So the wackestones-packstones occurring
mainly in Denée have been interpreted as the distal equivalents of the packstones-grainstones that dominated in
Salet.

5. Conclusions

The `black marble' of Denée, included within the Molignée Formation of Moliniacian age and
developed in the central part of the Dinant sedimentation area (corresponding to an intra-platform basin), is a
fossil conservation deposit and belongs more particularly to the `obrution deposits' of Seilacher et al. (1985).
The turbiditic sedimentation with smothering effect combined with deficient oxygenation of the bottom waters
favoured the exceptional preservation of the faunas (e.g. echinoderms, fishes) by inhibiting the development of
the necrophagous and saprophagous organisms during the deposition of the `black marble' facies sensu stricto.
The periodic confinement of the central part of the Dinant sedimentation area was induced by sea-level
fluctuations of low magnitude and took place during a third-order sequence characterized by a low sea-level,
namely the sequence 5 of Hance et al. (2001).

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130 YEARS AGO: THE DISCOVERY OF
THE BERNISSART IGUANODONS

Pascal GODEFROIT

Royal Belgian Institute of Natural Sciences, rue Vautier 29, 1 000 Bruxelles, Belgium,
Pascal.Godefroit@naturalsciences.be

Abstract: The discovery, in 1878, of more than twenty complete skeletons of the ornithopod dinosaur
Iguanodon, is one of the most important discoveries in the history of palaeontology. Here we shortly describe the
discovery, the excavation, the preparation, the exhibition and the study of these extraordinary skeletons.

KEYWORDS. Iguanodon, Bernissart, Lower Cretaceous, dinosaurs

1. Introduction

Bernissart is a former coal-mining village in western Belgium, situated less than a km from the Franco-
Belgian frontier. In 1878, the Sainte-Barbe Pit (Fig. 1) started to produce one of the greatest dinosaur discoveries
of all times: more than 20 complete articulated skeletons and several incomplete specimens of Iguanodon. These
were the first complete skeletons ever discovered and still remain one of the greatest accumulations of a single
taxon of dinosaur. This discovery was a cornerstone in the history of palaeontology: for the first time, it was
possible for the scientific community to realize how dinosaurs really looked like. Most of the specimens of
Iguanodon are now on display in the renovated Janlet Wing of the Royal Belgian Institute of Naturals Sciences
in Brussels. Nine of them are standing, mounted within an enormous glass cage. Many others have been left in
their original position, lying on their sides as found entombed in the coal mine. This astonishing array of
Iguanodon skeletons constitutes one of the most impressive displays of dinosaurs anywhere in the world.

Figure 1. The Sainte-Barbe pit and mine buildings at the end of the 19th Century, at the time when the
Bernissart Iguanodons were discovered.

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2. Before Bernissart

Around 1822, Mary Ann Mantell discovered large fossilized teeth while strolling in the Sussex
countryside in England. Her husband, the physician Dr. Gideon Mantell, was very intrigued by these fossils. He
described them and named them Iguanodon (`Iguana tooth'), because of their superficial resemblance to those of
living iguanas (Mantell, 1825). Iguanodon, the second representative of the group after Megalosaurus
(Buckland, 1824), was one the few chart members of the `Dinosauria', named by Richard Owen in 1842.

For 56 years, very little was known about Iguanodon and other dinosaurs. Mantell imagined these
antediluvian animals to be some kind of giant lizards with elongated bodies and sprawling limbs. In 1854, the
sculptor Waterhouse Hawkins erected a full-size reconstruction of Iguanodon for the Crystal Palace exhibition
centre in London as a rhinoceros-like heavy quadruped with a large spike on its nose.

The first partial dinosaur skeleton, named Hadrosaurus foulkii Leidy, 1858, was discovered in 1857 in
New Jersey. This skeleton was reconstructed in a bipedal gait at the Academy of Natural Sciences of
Philadelphia, but many questions were still left unanswered about the general appearance of dinosaurs.

3. The discovery of the Bernissart Iguanodons

In 1878, miners working in the the Sainte-Barbe Pit at Bernissart reported that the Luronne seam was
cut out, at 322 m depth, by what they called a `cran', a local name for pits formed by natural collapse through the
coal seams and filled with Lower Cretaceous clayey deposits normally located above the Coal Measures. The
miners had to traverse this `cran' as quickly as possible in order to rejoin the Luronne seam.

On February 28, two miners, J. Créteur and A Blanchard, found in the clays of the `Cran' what they
believed to be a tree trunk filled with gold. Many other specimens were collected by the miners in March. On
April 2, the local doctor L'Hoir and the mine manager G. Fagès concluded, that these strange objects were in
fact fossil bones filled with pyrite, the `fool's gold'. They rapidly sent fragments of fossil bones to several
Belgian specialists. P.J. Van Beneden, a zoologist from Leuven University, was the first to recognize among the
collected specimens teeth of the dinosaur Iguanodon (Van Beneden, 1878). On April 12, the management of the
colliery sent a telegram to E. Dupont, director of the Royal Museum of Natural History in Brussels, asking for
the services of L. De Pauw, a technician highly experienced in the restoration of fossils.

Figure 2. Drawing by G. Lavalette of a specimen of Iguanodon bernissartensis,
as discovered in the Sainte-Barbe pit at Bernissart.

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4. The excavation of the Bernissart Iguanodons

During three years, L. De Pauw and his team, composed by one museum warder, one moulder and nine
miners, actively excavated the `Iguanodon Cran' at Bernissart. In August 1878, an important earthquake blocked
the excavation team during two hours in the gallery 322 m below ground level. This gallery was subsequently
flooded in autumn, forcing the team to abandon their researches during several months. The excavations
restarted from May 1879 onwards. They were extended horizontally for 50m at the 322 m level. The miners also
encountered fossiliferous clays at a depth of 356 m, but the diameter of the `cran' was reduced at only 9m at this
level and the skeletons were consequently completely dislocated (De Pauw, 1902).

It was the first time that palaeontologist had the opportunity to collect such a wealth of fossils within a
single locality. More than twenty more or less complete skeletons of Iguanodon were found lying as they had
fallen, little disturbed by their burial (Fig. 2). Besides these dinosaurs, hundreds of fragments of plants, hundreds
of fishes, several crocodiles and tortoises, one amphibian, one fragment of insect, and one carnivorous dinosaur
phalanx were also discovered (Table 1).

Insects

Order Hemiptera

Hylaeoneura lignei Lameere & Severin, 1897
Fishes

Order Palaeonisciformes

Coccolepis macropterus Traquair, 1911

Order Pycnodontiformes

Lepidotes bernissartensis Traquair, 1911

Lepidotes brevifulcratus Traquair, 1911

Lepidotes arcuatus Traquair, 1911

Turbomesodon bernissartensis (Traquair, 1911)

Order Amiiformes

Callopterus insignis Traquair, 1911

Amiopsis dolloi Traquair, 1911

Amiopsis lata traquair, 1911

Notagogus parvus traquair, 1911

Order Pholidophoriformes

Pholidophorus obesus (Traquair, 1911)

Pleuropholis sp.

Order Gonorhynchiformes

Aethalionopsis robustus (Traquair, 1911)

Order Salmoniformes

Pattersonella formosa (Traquair, 1911)

Nybelinoides brevis (Traquair, 1911)

Order Elopiformes

Arratiaelops vectensis (Woodward, 1890)
Amphibian

Caudata incertae sedis

Hylaeobatrachus croyii Dollo, 1885
Turtles

Chitracephalus dumonii Dollo, 1885

Peltochelys duchasteli Dollo, 1885
Crocodiles

`Goniopholis simus Owen, 1878'

Bernissartia fagesiii Dollo, 1883
Dinosaurs

Iguanodon bernissartensis Boulenger, 1881

Iguanodon atherfieldensis Hooley, 1924

Theropoda indet

Table 1. Faunal list of the `Iguanodon cran' at Bernissart.

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The excavation method elaborated by L. De Pauw was so efficient that it is still used at the present time
during palaeontological excavations. Each Iguanodon skeleton was split into pieces that were coated with plaster
of Paris. After being sketched and catalogued, the blocks were carried to the surface. After three years of
excavations, about six hundred blocks, totalling more than 130 tonnes, were transported to Brussels in furniture
removal vans.

The excavations were stopped in 1881, because the expenses involved by this enterprise were
considered too high by the Belgian government. Members of the Parliament suggested that an Iguanodon
skeleton should be sold abroad in order to collect supplementary subsidies, but public outcry prevented this
transaction. From 1915, the German forces of occupation, under the initiative of the palaeontologist Otto Jaekel,
planned to start new excavations at Bernissart in order to send new Iguanodon skeletons in German natural
history museums. But the preliminary researches were interrupted in 1918 by the end of the First World War
(Roolf, 2004). After the war, further initiatives to start new excavations at Bernissart were immediately stopped
because of the absence of wish from the Belgian government to put up the money for such researches.

5. Preparation and mounting of the Iguanodon skeletons

After death, the Iguanodon carcasses were covered by clayey sediments and their decomposition
therefore developed in anoxic environment. In such conditions, sulphate-reducing bacteria were highly involved
in the putrefaction processes. The hydrogen sulphide, produced during the hydrolysis of the organic matter by
these bacteria, combined with the iron from the sediments and from the degradation of haemoglobin to form
pyrite, which was deposited in cavities within the bones. In contact with damp air, the pyrite oxidised to form a
salt, iron sulphate, or an iron oxide, limonite. Decomposition of both led to the disintegration of bone containing
them (Leduc, 2004). For that reason, the Iguanodon bones became extremely fragile when they were extracted
from the Bernissart pit.

Figure 3. Mounting of the first Iguanodon specimen in the St. George Chapel of the Nassau Palace.
Note, close to the Iguanodon's leg, the cassowary and wallaby skeletons used for comparison.

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Once they were arrived in Brussels, the Iguanodon blocks were stored in the Museum workshop,
housed in the St George Chapel of the Nassau Palace, now preserved as an exhibition hall in the Albert I Royal
Library. Between 1878 and 1905, the bones were impregnated with a carpenter's glue-based gelatine and the
pyrite was systematically curetted from the bones. Some vertebrae contained more than 1 kg of pyrite. The
remaining cavities were filled with `carton-pierre', a stable mixture of paper, glue and talc (De Pauw, 1902).

It was decided to mount the best preserved Iguanodon specimens in a lifelike gait. In 1882, the first
specimen was assembled and mounted by L. De Pauw and his team in the St. George Chapel. The bones were
suspended from scaffolding by ropes that could be adjusted so as to obtain the most lifelike position for the
complete skeleton, which was then supported by an iron framework (Fig. 3). This first mounted specimen was
publicly exhibited in 1883 in a glass cage constructed in the interior court of the Nassau Palace. But the Nassau
Palace Chapel quickly became too small for the storage, preparation, mounting and exhibition of these numerous
and bulky skeletons. In 1891, the Iguanodons and the Royal Museum of Natural History were transported to a
new home in the Leopold Park. In 1899, five specimens were mounted in a glass cage close to the entrance of the
museum. From 1902 onwards, the whole Bernissart exhibition was permanently installed in the newly-
constructed Janlet Wing of the Royal Museum of Natural History (Fig. 4).

Between 1933 and 1937, the Iguanodon skeletons were dismantled and treated, because thirty years of
changes in temperature and humidity had produced important damages. The bones were soaked in a mixture of
alcohol and shellac, a natural lacquer secreted by coccid insects. The specimens were installed into two large
glass cages, in order to stabilize the temperature and humidity of their environment.

Figure 4. Exhibition, in the beginning of the 20th Century, of the mounted Iguanodon
specimens in the Janlet Wing of the Royal Museum of Natural History.

From 2004 till 2007, the Janlet Wing of the Royal Belgian Institute of Natural Sciences was entirely
renovated. At this occasion, the Iguanodon skeletons were completely restored again. All the bones were
reinforced by a solution in acetone and alcohol of synthetic polyvinyl acetate (`Mowilith'). New glass cages
were constructed to protect the skeletons (Fig.5).

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Figure 5. The new presentation of the Bernissart Iguanodons.

6. The study of the Bernissart Iguanodons

Just after the discovery of the Bernissart Iguanodons, E. Dupont, then director of the Royal Museum of
Natural History, asked the young naturalist G. A. Boulenger to study these specimens. In 1881, Boulenger
presented his first results to the Belgian Academy of Sciences, Letters and Fine Arts: he described the anatomy
of the pelvis of these dinosaurs and proposed that the greater number of sacral vertebrae (six) in the Bernissart
form, as opposed to the five sacral vertebrae in the English species I. mantelli, merited the establishment of a
new species that he named Iguanodon bernissartensis. Unfortunately, this paper was refused publication,
although a brief highly critical review of Boulenger's paper was published by Van Beneden (1881). The latter
claimed that the Bernissart Iguanodons belonged to Iguanodon mantelli, already described from disarticulated
specimens discovered in England. Shortly afterwards in 1881, Boulenger accepted a post at the British Museum
(Natural History) and in 1882 study of the Bernissart Iguanodons was entrusted to L. Dollo. Between 1882 and
1923, Dollo published many preliminary notes on the Bernissart fauna and, especially, on Iguanodon (see
bibliographical list in Norman, 1980). He distinguished two species at Bernissart: most of the specimens belong
to the larger form I. bernissartensis, the new species named by Boulenger, although a single complete individual
represents the smaller and slender I. mantelli. While studying in detail several parts of the Iguanodon skeleton,
Dollo was adopting a forensic approach to understanding these fossils. He developed a new style of
palaeontology that became known as palaeobiology: palaeontology can be expanded to investigate the biology,
and by implication the ecology and the behaviour of extinct creatures. Dollo's final contribution to the
Iguanodon story was published in 1923 as a synthetic study, to honour the centenary of Mantell's original paper.
He identified Iguanodon as an ecological equivalent of the giraffe. Its kangaroo-like posture enabled it to reach
high into the trees to gather its fodder, which it was able to draw into its mouth by using a long, muscular tongue.
The sharp beak was used to nip off tough stems, while the teeth served to pulp the food before it was swallowed.
This image of Iguanodon as a gigantic kangaroo-style creature, as depicted by Dollo, has become iconic during
more than 60 years and was reinforced by the distribution of full-sized replicas of mounted skeletons of
Iguanodon from Brussels to many of the great museums around the world (Norman, 2005).

In 1980, the British palaeontologist D. Norman published a monographic study of Iguanodon
bernissartensis. Functional analysis of the skeleton indicated that the vertebral column, stiffened by a network of
ossified tendons, was held more or less horizontal while the animal was walking or running. Norman also
believed that I. bernissartensis was mainly a quadrupedal animal. The structure of the pectoral girdle, the ratios
of the fore-and hindlimb lengths, the strongly fused carpal bones, and the presence of hoof-like unguals on the

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middle three digits of the hand suggest that the adult of I. bernissartensis spent most of its time in a quadrupedal
posture, though juveniles had a predominantly bipedal mode of life.

In 1986, Norman concluded that the small species from Bernissart belongs to Iguanodon atherfieldensis
Hooley, 1925, a species previously described from the Wealden Beds of the Isle of Wight.

Figure 6. Reconstruction by C-T scanning of the skull and brain of Iguanodon bernissartensis.

Although the Bernissart Iguanodons were discovered 130 years ago, these fantastic creatures have not
revealed all their secrets yet. At the occasion of the last dismounting operation of the Iguanodon specimens, the
best preserved skulls have been investigated by CT-scanning at Gasthuisberg Hospital in Leuven. It is now
possible to study inaccessible areas inside the skull without damaging these fragile fossils. Fig. 6 is a
reconstruction, after C-T scanning, of the brain cavity of these animals, dead some 130 million years ago.

7. References

BUCKLAND, W., 1824. Notice on Megalosaurus or great fossil lizard of Stonesfield Transactions of the Geological Society of London, 21:
390-397.
DE PAUW, L.F., 1902. Notes sur les fouilles du charbonnage de Bernissart. Découverte, solidification et montage des Iguanodons. Imprim.
photo-litho, JH. & P. Jumpertz, 150 av.d'Auderghem, Bruxelles, Belgium.
LEDUC, T., 2004. La diagenèse des ossements fossiles: étude bibliographique. Mémoire de D.E.A. non publié, Université de
Liège, Belgium.
LEIDY, J., 1858. Hadrosaurus and its discovery. Proceedings of the Academy of Natural Sciences, Philadelphia: 213-218.
MANTELL, G.A., 1825. Notice on the Iguanodon, a newly discovered fossil reptile, from the sandstone of Tilgate Forest, in Sussex.
Philosophical Transactions of the Royal Society, 115: 179-186.
MARTIN, F. & BULTYNCK, P., 1990. The Iguanodons of Bernissart. Institut royal des Sciences naturelles de Belgique, Bruxelles,
Belgium.
NORMAN, D.B., 1980. On the ornithischian dinosaur Iguanodon bernissartensis of Bernissart (Belgium). Mémoires de l'Institut royal des
Sciences naturelles de Belgique 178: 1-103.
NORMAN, D.B. 1986. On the anatomy of Iguanodon atherfieldensis (Ornithischia, Ornithopoda). Bulletin de l'Institut royal des Sciences
naturelles de Belgique, Sciences de la Terre 56: 281-372.
NORMAN, D.B., 2005. Dinosaurs. A very short introduction. Oxford University Press, Oxford, U.K.
OWEN, R., 1842. Report on British fossil reptiles, part II. Report of the Eleventh Meeting of the British Association for the Advancement of
Science, held at Plymouth, July 1841, 66-204. London.
ROOLF, C., 2004. Dinosaurier-Skelette als Kriegziel: Kulturgutraubplanungen, Besatzungpolitik und die deutsche Paläontologie im Ersten
Weltkrieg. Berichte zur Wissenschaftgeschichte, 27: 5-26.
VAN BENEDEN, P.-J., 1878. Découverte de reptiles gigantesques dans le charbonnage de Bernissart, près de Péruwelz. Bulletin de
l'Académie royale de Belgique, 2ème série, 45: 578.
VAN BENEDEN, P.-J., 1881. Sur l'arc pelvien chez les dinosauriens de Bernissart. Bulletin de l'Académie royale belge, classe des Sciences,
3 : 600-608.

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THE LOWER CRETACEOUS WEALDEN FACIES OF THE
HAUTRAGE QUARRY (MONS BASIN, BELGIUM)

Johan YANS 1, Paul SPAGNA 2, Thomas GERARDS 3, Philippe GERRIENNE 3, Jean
DEJAX 4, Bernard GOMEZ 5, Christian DUPUIS 2, Jean-Marc BAELE 2, François
BARBIER 1, Stéphane PIRSON 6

1 FUNDP, UCL-Namur, 61 rue de Bruxelles, B-5000 Namur, Belgium. johan.yans@fundp.ac.be,
francois.barbier@fundp.ac.be
2 FPMS, Service de géologie fondamentale et appliquée, 9 rue de Houdain, B-7000 Mons, Belgium.
Paul.Spagna@fpms.ac.be, Christian.Dupuis@fpms.ac.be, Jean-Marc.Baele@fpms.ac.be
3 Université de Liège, Paléobotanique, Paléopalynologie et Micropaléontologie, B18, Sart-Tilman, 4000 Liège.
tgerards@hotmail.com, p.gerrienne@ulg.ac.be
4 MNHN, USM 0203, CNRS UMR 5143 (Paléodiversité et paléoenvironnements), Département Histoire de la
Terre, case postale 38, 57 rue Cuvier, 75231 Paris Cedex 05, France, dejax@mnhn.fr
5 Université Lyon 1 (Claude Bernard), CNRS-UMR 5125 Paléoenvironnements et Paléobiosphère (PEPS) 43
Bd du 11 Novembre 1918, 69622 Villeurbanne Cedex, France, bernard.gomez@univ-lyon1.fr
6 Institut royal des Sciences naturelles de Belgique, Département de Paléontologie, 29 rue Vautier, B-1000
Bruxelles, Belgium. stef.pirson@skynet.be

Keywords: Hautrage, Mons basin, Belgium, Wealden facies, Early Cretaceous

1. Introduction to the geology of Belgium

The Hautrage quarry is located in the Western part of Belgium. Despite the small size of its territory
(ca. 30.500 km²), Belgium shows a rich geology, encompassing various lithologies and covering a large part of
the geological timescale, from Early Palaeozoic to Holocene. Figure 1 focuses on the geology of Belgium.

Figure 1. Simplified geological map of Belgium.

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Excepting the Quaternary cover, rocks outcropping in Belgium can broadly be divided into two large
areas (Fourmarier, 1954; Robaszynski & Dupuis, 1983).
The northern part of the country mainly exhibits Cenozoic deposits consisting in predominantly marine
and unconsolidated sediments (Vandenberghe et al., 1998). They may reach a thickness of several hundreds of
meters. Sediment accumulation in this area resulted from relative sea-level fluctuations and migration of the sea
to the North/North-West.
In southern Belgium consolidated Palaeozoic rocks dominate over large areas. These rocks underwent
strong deformations at the end of Carboniferous. Mesozoic deposits are poorly represented due to dominant
continental conditions during this period. Marine sediments from Triassic and Jurassic only crop out in southern
Belgium (Belgian Lorraine; Boulvain et al., 2001). Rocks from that period are also known from boreholes in the
Campine Basin (Dusar et al., 2001). Continental deposits of the Lower Cretaceous are only preserved in the
Mons basin, in which the famous Bernissart Iguanodon were found in 1878-1881 (Robaszynski et al., 2001).
More or less extended marine deposits formed during Upper Cretaceous due to transgressive pulses. The
Cretaceous sea progressively flooded a large part of Belgium (Robaszynski et al., 2001). Cretaceous strata
mainly crop out in the Liège-Maastricht area and in the Mons basin. In the southern part of Belgium, Cenozoic
deposits are rather poorly represented, except in the Mons basin area.
Almost all these rocks are of sedimentary origin (Bultynck & Dejonghe, 2001). Occurrences of magmatic rocks
are rather rare (Denaeyer & Mortelmans, 1954) and metamorphic rocks are restricted to the Ardenne
Anticlinorium and the Brabant Massif (Fielitz & Mansy, 1999).

2. The Mons basin

The Mons basin (MB) is located in the western part of Belgium, in the Hainaut Province (Figure 2). It is
connected to the Paris Basin to the West although the sedimentary records are significantly different in both
basins. The MB may be considered as a gentle "syncline" developed on a folded and faulted Devonian-
Carboniferous basement (Fig. 6A), and filled with Cretaceous-Cenozoic sediments (Robaszynski et al., 2001).
The MB is limited by the maximal extension of the Turonian deposits.

Figure 2. Schematic geological map of the Mons basin.

2.1. Lithostratigraphy

The deposition of the sediments in the MB is the result of relative sea-level fluctuations, due to
eustatism, subsidence and sedimentary supplies (Figure 3).

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Figure 3. A. North-South section in the Mons basin; vertical amplitude = x 10. B. Simplified lithostratigraphy of
the Mons basin (from Marlière, 1970).

The oldest sediments of the MB are the "Wealden facies" recognized in three geological contexts:
- in kilometric outcrops ("pockets") or weakly buried sediments in the northern part of the MB, from Hautrage to
La Louvière (Fig. 2; Marlière, 1946) Hautrage Clays Formation, Baudour Clays Formation and Saint-Pierre
Clays Formation;
- in filling several natural pits (also called "Cran") developed through the basement (for example at Bernissart);

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- as white sands and sandstones containing lignite and glauconitic material in the eastern part of the MB (=
"Strepy Formation" or "Cénomanien à faciès Wealdien" sensu Gulinck, 1974).
In the MB, the "Wealden facies" are locally covered by calcirudites, sandstones and conglomerates of
the Albian-Cenomanian Haine Group (locally called "Meule", Marlière, 1946). Rich in glauconite and
ferriferous illite, these Albian-Cenomanian deposits correspond to neritic sedimentation and can reach 190
meters of thickness in local depressions for example at Bernissart - where the subsidence rate is higher than in
adjacent areas (Cornet, 1927). These depressions, called "Cuves", are scattered throughout the basin.
The latest Cenomanian period characterizes the base of a large sea-level rise (probably due to eustatism) with the
glauconiferous marls rich in Actinocamax plenus. Marls mainly dominate in the overlying Turonian sediments
("Dièves"), showing frequent lateral variation of thickness. The Upper Turonian - Coniacian sedimentation
probably recorded the eustatic regression / transgression event which is known world-wide during this period
(Hacq et al., 1987). Upper Turonian deposits consist of 5 to 40 meters of siliceous marls (Chailles de Ville-
Pommeroeul) overlain by cherty limestone (Silex d'Hautrage) that exhibit increasing lateral changes with time
and are capped by hardgrounds. On top of the latter, a 0.5 to 2m-thick bed of highly glauconitic sediments
("Craie" de Maisières, probably Coniacian in age) mark the lower, transgressive sequence of the white chalk
deposits.
Chalks are widespread in the MB during the Coniacian, Santonian, Campanian and Maastrichtian
stages. The chalk series is locally covered by the phosphatic "Craie de Ciply" and by the "Tuffeau de Saint-
Symphorien". During Late Maastrichtian, the MB experienced a (eustatic?) sea-level fall with condensed beds
(hardgrounds) and hiatuses. The famous Cretaceous-Palaeogene boundary is thus not recorded in the MB, at
least in known sections.
The Cenozoic sedimentation starts with the "Dano-Montian" and "Montian", Danian (-Selandian) in
age. It partly corresponds to continental deposits with local mammal fossils like those excavated at Hainin,
which are considered as a key-point for the stratigraphy of mammals in Europe (Folie et al., 2005). The latest
Palaeocene consists in argillaceous, glauconiferous and locally carbonaceous Thanetian sands. During the
Palaeocene-Eocene interval, continental conditions prevailed in the MB as shown by the occurrence of fluviatile
deposits, locally containing terrestrial fauna (like in Erquelinnes: Rutot, 1881). Continental conditions are further
attested by a large meteoric weathering of the latest Thanetian marine sands and the fluviatile sands, resulting in
widespread quartzitic concretions (Dupuis et al., 1997). The early Eocene is characterized by Ypresian sandy
clays. Coarse Lutetian sand is located only in the eastern part of the MB.

2.2. Subsidence

As demonstrated in the Saint-Ghislain borehole (Delmer, 1972), high subsidence rates in the Mons area
are recorded since Palaeozoic times. Kilometric-thick Upper Devonian and Carboniferous series accumulated in
the EW-elongated basin called "sillon borain" by Michot (1980). Abnormal but of lower order of magnitude
subsidence rates are still observed through the Meso-Cenozoic, with sedimentary sequences of ca. 200 m-thick in
average. The most striking feature of subsidence in the MB is depocentre shift (Marlière, 1970). The thickness
maxima for each sedimentary formation are not superimposed, showing that the location of areas of highest
subsidence rate changed with time. The thickness maxima of the different formations in the Cenomanian-
Lutetian sequence give 800 m when summed together. This demonstrates the significant impact of depocentre
shift on sedimentary record in the MB. As a result of subsidence and except along fault-flexure zones where
layers locally dip as 60°, bedding dips are rather small (about 10°; Angelier et al., 2006). The existence of
evaporite dissolution in the deep Devono-Dinantian basement and its impact on the deformation of the overlying
strata have been suggested to explain the depocentre shift (Delmer, 1972; Dupuis & Vandycke, 1989). However
the very well oriented fault system in the MB may also be partly due to the regional crustal activity (Vandycke et
al., 1991; Vandycke, 2002; Spagna et al., 2007). "Cuves", "pockets" and "Crans" are local high subsidence
areas, with infillings of Lower Cretaceous and Lower-Middle Cenomanian deposits.

3. The Hautrage quarry

3.1. Localization and geological setting

The Danube-Bouchon quarry is localized in a "pocket", called the "Hautrage pocket" (Saint-Ghislain
entity) at about 20 kilometers north-west of Mons. The wealden clays and sands trapped in this pocket belong to
the Hautrage Clays Formation. They overlie the Namurian weathered basement, and are recovered by
Cenomanian-Turonian marine sediments. In Hautrage, the wealden facies are deposited in a continental system.
It can be linked to a major gully that exists between the south-east English area and the future Paris Basin (Thiry
et al., 2006). The relative low sea-level of this period probably allows this gully to collect the products of the
surrounding Armorican Massif in the south west, and London-Brabant Massif in the north east.

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The CBR-Heidelbergcement industry mines and uses the clays for their high aluminate and silicate
contents. During prospective campaigns several boreholes have been realized, sampled and analyzed, in order to
decipher the clays and sands qualities for their future applications. To complete the data sets, many cross
sections have been described and analyzed during and/or after the successive annual exploitation campaigns.
Recently, a modeling has been developed to improve the mining (Figure 4).

Figure 4. Modeling of the Hautrage quarry.

Figure 5. North-South lithological cross-section through the Danube-Bouchon quarry of Hautrage (from Spagna
2005).

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Figure 6. General succession of the Hautrage Clays Formation (from Yans, 2007; Spagna et al., 2008).

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3.2. Sedimentology

The Danube-Bouchon quarry cuts clayey and sandy sediments rich in organic matter (root traces,
lignite, wood fragments, ...), sideritic nodules and pyrites. The sediments are clearly stratified and east-west
oriented, global dipping south with 15 to 25° dip, as shown by different key beds all along the series (Figure 5).
As shown in the synthetic log at Figure 6, the Hautrage succession of 235 meters of wealden sediments can
be divided into 3 parts (Yans, 2007; Spagna et al., 2008):
a) dominance of red "clays" (however all the analyzed samples contained around 60-70% of micrometric to
millimetric grains of quartz, making them belong to the "clayey silt and sands" or "silty and sandy clays"
categories, but we will use the "clays" term for more usability) rich in weathered siderite at the bottom
(Units A to D)
b) alternations of black, grey, white and bluish clays containing various quantities of sands, wood fragments,
pyritic and sideritic nodules (Units E to I)
c) dominance of sandy (to conglomeratic) sediments rich in pyrite and wood fragments at the top (J and K
Units).
Traces of paleosoils can be found in the Hautrage wealden facies, especially in the lower part of the deposit:
- the red color (and marmorosis) of the clays found at the bottom of the series is related to a weathering
(pedogenic oxidizing) of their primitive sideritic contain;
- "black pebble" have been observed together with illuviation figures, rhizolite and crackings, all of which
indicate a poor maturation of the soil;
- traces of roots are common, sometimes trapped in sideritic nodules.
All these elements indicate the exundation and stability of the sediments during relatively long periods. An in
situ tree stump has also been found within lightly variegated clays.
Channels are present at different levels in the quarry. The massive sandy outcrop of the southern quarry
front (J Unit) is of fluvial origin. Its pebble contain gives an idea of the level of energy that could have been
developed at that time. During intense rising events, channels were overflowed, and clayey sediments, due to the
lost of energy, could then settling the plain out. The result of these floods may have been a relatively humid
environment adapted for the development of a swampy vegetal cover.

3.3. Mineralogy

Quartz- is the predominant mineralogical phase of the whole series. The clayey fraction (< 2 µm) is
divided in three main compounds: kaolinite, smectite (I-S mixed-layers) and illite. The mineralogical log of
Figure 6 shows the ratio of the other mixed-layer minerals, and illustrates a very clear trend of raising content of
the chlorite-smectite mixed-layer phase to the base of the succession. Seven mineralogical units can be defined
in the whole series, rather unlinked to the lithological one (Yans, 2007; Spagna et al., 2008).

3.4. Dating

In the last decade, palynological studies of new boreholes and new sections allow to precise the age of
the "Wealden facies" of the MB (Dejax et al., 2008). The palynomorphs of the "Wealden facies" from the
western part of the MB - i.e. Hautrage Clays Formation, Baudour Clays Formation and Sainte-Barbe Formation
are continental only. The botanical affinities of the palynological taxa are mainly ferns, gymnosperms and
freshwater algae. The stratigraphy of the angiospermous pollen grains suggests a middle Barremian to earliest
Aptian age for the Hautrage Clays Formation, Sainte-Barbe Clays Formation and Baudour Clays Formation
(Yans et al., 2005a, 2005b, 2006; Dejax et al., 2007a, 2007b).
On the other hand, the "Wealden facies" of the eastern part of the MB are Late Albian in age and do
contain dinoflagellates suggesting marine influences (Yans et al., 2007). The "Cénomanien à faciès wealdien" or
"Strépy Formation" is Turonian in age (Yans, 2007).
The Wealden facies may have supplied the filling of regional endokarsts during Late Cretaceous to
Early Cenozoic (Quinif et al., 2006).

3.5. Leaves, reproductive organs and wood

One of the most impressive features of the Hautrage Clays Formation is its content in meso- and
megafossil plant remains (including fragments of wood; Gomez et al., 2008; Gerards et al., 2008). They are
preserved as fusains (charcoals), conifer leafy axes (Cheirolepidiaceae Frenelopsis and Taxodiaceae cf.
Sphenolepis) bearing cuticles as well as inner anatomy, female cone scales (Cheirolepidiaceae Alvinia) and
cones. The lignite bed includes a much dissimilar assemblage consisting of amber, more abundant fusain
fragments, sterile and fertile fern pinnules and pinnae (Osmundaceae Cladophlebis, Weichseliaceae Weichselia,

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Matoniaceae Phlebopteris), conifer leaves and twigs (Miroviaceae Mirovia and Taxodiaceae cf. Sphenolepis),
reproductive organs and seeds.
The wood assemblage includes specimens of Podocarpoxylon, Taxodioxylon, Thujoxylon, cf.
Sequoioxylon, Brachyoxylon, three different Pinaceae, and 2 new genera. Tree ring analysis confirms the
palaeogeographical position (around 35°N) of the Mons basin during the Early Cretaceous. The composition of
the wood assemblage and the tree ring analysis suggest a humid warm-temperate climate. Taxodiaceae,
Podocarpaceae and various ferns thrived in the floodplain, probably in swampy environments. Mangroves with
Cheirolepidiaceae possibly existed downstream. Pinaceae and Cupressaceae lived in the surroundings.

4. Acknowledgements

Thanks are due to the Lebailly S.A. in Hautrage, to Wanty S.A. and CBR-Heidelbergcement industry
for their logistic cooperation.

5. References

ANGELIER, J., VANDYCKE, S., BERGERAT, F., GAVIGLIO, P., SCHROEDER, C. & COULON, M., 2006. Can belemnite distribution
reveal pressure-solution processes along faults? A case study in the chalk of the Mons Basin, Belgium. Journal of Structural
Geology, 28: 64-82.
BOULVAIN, F., BELANGER, I., DELSATE, D., DOSQUET, D., GHYSEL, P., GODEFROIT, P., LALOUX, M., ROCHE, M.,
TEERLYNCK, H. & THOREZ, J., 2001. New lithostratigraphical, sedimentological, mineralogical and palaeontological data on
the Mesozoic of Belgian Lorraine: a progress report. Geologica Belgica, 3: 3-33.
BULTYNCK, P. & DEJONGHE, L. (éd.), 2001. Guide to a revised lithostratigraphic scale of Belgium, Geologica Belgica, no special, 4, 168
p.
CORNET, J., 1927. L'époque wealdienne dans le Hainaut. Annales Société géologique de Belgique, L: 89-103.
DEJAX, J., PONS, D. & YANS, J., 2007a. Palynology of the dinosaur-bearing Wealden facies sediments in the natural pit of Bernissart
(Belgium). Review of Palaeobotany and Palynology, 144: 25-38.
DEJAX, J., DUMAX, E., DAMBLON, F. & YANS, J., 2007b. Palynology of Baudour Clays Formation (Mons Basin, Belgium): correlation
within
the
"stratotypic"
Wealden.
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Geology
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DEJAX J., PONS D. & YANS J. (2008). Palynology of the Wealden facies from Hautrage quarry (Mons Basin, Belgium). Memoirs of the
Geological Survey 55, 45-52.
DELMER, A., 1972. Origine du Bassin crétacique de la Vallée de la Haine. Bruxelles, Service géologique de Belgique (Professional paper,
5), 13 p.
DENAEYER, M.-E. & MORTELMANS, G., 1954. Les roches éruptives. In: FOURMARIER P. (éd.), Prodrome d'une description géologique
de la Belgique, Liège, Société géologique de Belgique, 745-792.
DUPUIS, C., EGGERMONT, B. & HENNEBERT, M., 1997. Evidences of tectonic movements during the Paleocene-Eocene interval (P/E,
NP8/NP11) in the Tournai area (Mélantois) and neighbouring regions (NW Paris Basin, Mons Basin). Aardkundige
Mededelingen, 8: 73-74.
DUPUIS, C. & VANDYCKE, S., 1989. Tectonique et karstification profonde: un modèle de subsidence original pour le Bassin de Mons.
Annales de la Société géologique de Belgique, 112, 2: 479-487.
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