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Fossilium Catalogus I:Animalia FOSSILIUM CATALOGUS I:ANIMALIA Edited by Wolfgang RIEGRAF Pars 144 Lower Cretaceous Ammonites III Bochianitoidea, Protancyloceratoidea, Ancyloceratoidea, Ptychoceratoidea by Jaap KLEIN, Robert BUSNARDO, Miguel COMPANY, Gerard DELANOY, Mikheil KAKABADZE, Stephane REBOULET, Pierre ROPOLO, Zdenek VAsicEK & Jean VERMEULEN IB Backhuys Publishers, Leiden, 2007 Fossilium Catalogus I: Animalia Edited by Wolfgang RrEGRAF Pars 144 Lower Cretaceous Ammonites III Bochianitoidea, Protancyloceratoidea, Ancyloceratoidea, Ptychoceratoidea by Jaap KLEIN, Robert BUSNARDO, Miguel COMPANY, Gerard DELANOY, Mikheil KAKABADZE, Stephane REBOULET, Pierre ROPOLO, Zdenek VAsicEK & Jean VERMEULEN IB Backhuys Publishers, Leiden, 2007 Copyright 2007 Backhuys Publishers All rights reserved. No part ofthis book may be translated or reproduced in any form by print, photoprint, micro­ film, or any other means without prior written permission ofthe publisher. Addresses of the authors: Robert BIJSNARDO: Dcpartcmcnt des Sciences de la Terre, Universite Claude Bcrnard Lyon 1,43 Boulevard du I1 Novcmbrc, 1'-69622 Villeurbanne cedex, France. Miguel COMPANY: Departamento de Estratigrafia y Paleontologfa, Facultad de Ciencias, Universidad de Granada, 18002 Granada, Spain. Gerard DELANOY: Universite de Nice-Sophia Antipolis, Departcmcnt des Sciences de la Terre, 28 Avenue Valrose 1'-06100, Nice cedex, France. Mikhcil KAKABADZE: A. Djanclidze Geological Institute, M. Alexidze str. 1/9,0193, Tbilisi, Republic of Georgia. Jaap Kl.EIN: Nationaal Natuurhistorisch Museum, Demmerik 12, NL-3645 EC Vinkeveen, The Netherlands. Pierre Roroi.o: Centre de Paleontologic-Scdimcntologic de I'Universite de Provence, Place Victor Hugo 1'-13331, Marseille, France & Centre d'etudes meditcrraneennes, Barremc, France. Stephanc REBOUl.ET: UFR des Sciences de La Terre, CNRS-UMR-5125 Paleoenvironnements et Paleobiosphere, Univcrsite Lyon 1. Batiment Geode, 2 rue Raphael Dubois, Campus de la Doua, 1'-69622 Villeurbanne cedex, France. Contribution to UMR5125-07.024. Zdcnck VAsiCEK: Institute ofGeonics (Academy of Sciences ofthe Czech Republie), Studentska 1768, CZ-708 00 Ostrava-Poruba, Czech Republic. Jean VERMEUl.EN: Grand rue, r-04330 Barrcme, France. Editor's address: Dr. Wolfgang RrEGRAF Briiggefeldweg 31, 0-48161 Miinster, Germany phone +49 0251 866 870 ISBN/EAN 978-90-5782-190-5 Cover: Ancvloceras Puzosianus D'ORBIGNY, 1842 Published by: Backhuys Publishers, P.O. Box 321, NL-2300 AH Lcidcn, The Netherlands Preface & acknowledgements During the last decades a lot of research on the heteromorphs of the Lower Cretaceous has been per­ formed: detailed stratigraphic research ofValanginian, Hauterivian, Barremian and Aptian key sections in SE France, in the rest ofEurope, in America, and in Africa, precise revision ofclassical ammonite collections and significant work on their morphology and evolution. The heterornorphs are difficult to study and to interpret because they are a complex group ofam­ monites. Gerard DELANOY wrote about his field ofresearch: "There arc many small heteromorphic am­ monites with tuberculated venter in the Upper Barremian and they arc difficult to integrate in a phylogenetic scheme because we do not always have all the elements. Heteromorphic ammonites have only a small number of ornamental and morphological characters and they "play" always with the same: uni-bi-trituberculate ribs, non tuberculate ribs, vigorous ribbing on the body chamber or not, coiled or uncoiled shells with all the intermediate forms. All these characters can change very quickly during their range". Thereto we can add that a lot of heterornorph species are based on very frag­ mentary material ofunknown stratigraphical origin, that too many species are based on a single spec­ imen, that isolated findings from outside the Tethys are difficult to integrate in a classification and that a thorough revision ofthe Lower Cretaceous heteromorph fauna ofGermany is unfortunately not yet done. So I am very pleased that nearly all specialists that are active in this research were willing and could find time to work together on this part of the Fossilium Catalogus. In his thesis Stephane REBOULET made it plausible that the superfamily Ancyloceratoidea is prob­ ably not monophyletic but has a multiple origin in the Neocomitidae. He described possible ancestors in the Valanginian and Lower Hauterivian. For decades Jean VERMEULEN studied the stratigraphy and ammonite distribution in the Upper Hauterivian and Lower Barremian ofSE France, in the first years working on the Pulchelliidae but in the last years also on heteromorphs. His recent publications about a new phylogenetic classification ofthe Lower Cretaceous hcteromorphs were the start of heated debate. Gerard DELANOY wrote a lot of papers and books on ammonite stratigraphy, especially on the Upper Barremian and around the Barrernian-Aptian boundary in SE France. Furthermore he pub­ lished papers on the classification of, and the evolution ofcoiling in, heteromorphs. The ammonite stratigraphy around the top ofthe Barremian and the base of the Aptian in La Be­ doule was studied by Pierre ROPoLo. He published significant papers on the evolution and variation ofcoiling and on the dimorphism of heterornorphs. In Spain Miguel COMPANY and his colleagues from Granada were the first who investigated the Lower Cretaceous of the Bctic Cordillera of Spain in detail, especially the ammonites and their dis­ tribution around the Berriasian-Valanginian boundary, in the Valanginian, Barremian, Aptian, and around the Barremian-Aptian boundary. The achievement of Zdenck VAsicEK is a long list of publications on the Lower Cretaceous am­ monites and stratigraphy ofthe Carpathians ofthe Czech and Slovak Republics, Upper Austria and the Northern Calcareous Alps. He revised the ammonites described by UHLIG ofthe "Ieschener & Werns­ dorfer Schichtcri'. Recently Robert BUSNARDO revised most ofthe Lower Cretaceous ammonites described by Alcide D'ORBIGNY in his 'Paleontologie francaise, Terrains Crctaces. Cephalopodes'. He researched together with JEAN CHAROLLAIS, MARC WEIDMANN & BERNARD CLAVEL the Early Cretaceous ofLa Veveyse dc Chatel in Switzerland. They could place most of the ammonites described by OOSTER, SARASIN & SClIONDELMAYER and PICTET & DE LORloL in a strati graphical context. Mikheil KAKABADZE is well known from his work on the heteromorphs of the Caucasus, Crimea and Colombia. Besides he wrote articles on the mode of life, the classification, and on the intraspe­ cific and intrageneric variabilities ofthe heteromorph ammonites. v Although we all disagree about a lot in this Fossilium Catalogus, however, we consider our result as a starting point for further research on the classification ofthe Lower Cretaceous heteromorphs. In addition to the acknowledgements given in Part I and II ofLower Cretaceous Ammonites we are obliged to Dr. Francisco NULLO (Servicio Geol6gico Nacional) from Argentina, Dr. Coral STANLEY (South Australian Parliamentary Library) from Australia, Dr. Michal KROBICKI (University ofMining and Metallurgy, Krak6w) from Poland, Dr. Mikhail ROGOV (Geological Institute ofthe Russian Acad­ emy ofSciences) from Russia and Dr. Peter RODDA (California Academy ofSciences) from the United States for helping us with literature. Special thanks to Prof. Dr. Fabrizio CECCA (Universite Pierre et Marie Curie, Paris) and Prof. Dr. Gerard THOMEL (St. Blaise) for providing us with additional infor­ mation. Vinkeveen, in summer 2007 Jaap KLEIN vi Contents INTRODUCTION I Superfamily BOCHIANITOIDEA SPATH, I922 2 FamilyBOCHIANITIDAE Spath, 1922 2 Superfamily PROTANCYLOCERATOIDEA BREISTROFFER, 1947 13 Family PROTANCYLOCERATIDAE Breistroffer, I947 13 FamilyAEGOCRIOCERATIDAE Venneulen, 2006 21 Superfamily ANCYLOCERATOIDEA GILL, I871 31 FamilyHIMANTOCERATIDAE Dimitrova, 1970 31 Family CRIOCERATITIDAE Gill, 1871 34 Family EMERICICERATIDAE Vermeulen, 2004 67 FamilyACRIOCERATIDAE Vermeulen, 2004 94 FamilyANCYLOCERATIDAE Gill, 1871 118 FamilyHETEROCERATIDAE Spath, 1922 174 FamilyHEMIHOPLITIDAE Spath, 1924 213 Family LEPTOCERATOIDIDAE Thieuloy, 1966 230 Family LEPTOCERATOIDIDAE genus incertum 244 FamilyLABECERATIDAE Spath, 1925 245 Superfamily ANCYLOCERATOIDEA familia incerta 260 Superfamily ANCYLOCERATOIDEA genus incertum 260 Superfamily PTYCHOCERATOIDEA GILL, 1871 264 FamilyPTYCHOCERATIDAE Gill, 1871 264 FamilyHAMULINIDAE Gill, 1871 271 '?Family MEGACRIOCERATIDAE Vermeulen, 2006 285 Family MACROSCAPHITIDAE Hyatt, 1900 289 References 305 INDEX 355 VII Introduction The classification in this part ofthe Fossilium Catalogus differs from that ofWRIGHTet al., 1996. Three new superfamilies are introduced: the Bochianitoidea SPATH, 1922, the Protancyloceratoidea BREISTROFFER, 1947 and the Ptychoceratoidea GILL, 1871 in which several families and genera are listed that WRIGHT included in the superfamily Ancylocerataceae (Ancyloceratoidea) GILL, 1871. The superfamilies Bochianitoidea and Protancyloceratoidea are excluded from this superfamily because representatives probably derived from Late Jurassic perisphinctids and have no phylogenetic lineages with the Ancyloceratoidea. The origin ofthe Ptychoceratoidea is yet unclear. VERMEULEN, 2006 classified this superfamily in the Turrilitina BESNOSOV & MIKHAILOVA, 1983. The subfamily Helicancylinae HYATT, 1894 is treated as being invalid. WRIGHT himselfregards this not as a natural subfamily but a collection ofsmall forms and microconchs of various late Crioceratitinae and Ancyloceratinae. Following VERMEULEN, 2004 and 2006 we tentatively
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