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Home , Actinocerida, Ammonitina, Aulacocerida, Goniatitina, La Caja Formation, Placenticeratidae

Advancing Research on Living and Development and Evolution Fonn, Construction, and Function Taphonomy, Palaeoecology, Palaeobiogeography, , and Basin Analysis Advaociog Research 00 Living and Fossil Cephalopods

Development and Evolution Form, Construction, and Function Taphonomy, Palaeoecology, Palaeobiogeography, Biostratigraphy, and Basin Analysis

Edited by

Federico 016riz and Francisco J. Rodriguez-Tovar University of Granada Granada. Spain

Springer Science+Business Media, LLC Library of Congress Cataloging in Publication Data Advancing research on living and fossil cephalopods: development and evolution: form, con• struction, and function: taphonomy, palaeoecology, palaeobiogeography, biostratigraphy, and basin analysis I edited by Federico Ol6riz and Francisco J. Rodriguez-Thvar. p. cm. "Proceedings of the IV International Symposium on Cephalopods: Present and Past, held July 14-18, 1996, in Granada, Spain"-T.p. verso. Inc1udes bibliographical references and index. ISBN 978-1-4613-7193-9 ISBN 978-1-4615-4837-9 (eBook) DOI 10.1007/978-1-4615-4837-9 1. Cephalopoda, Fossil-Congresses. 2. Cephalopoda-Congresses. 1. 0l6riz Sâez, Fed• erico. II. Rodriguez-Thvar, Francisco J. III. International Symposium on Cephalopods: Pre• sent and Past (4th: 1996: Granada, Spain) QE806.A38 1998 98-41749 564'.5-dc21 CIP

Proceedings of the IV International Symposium on Cephalopods: Present and Past, held July 14 - 18, 1996, in Granada, Spain

ISBN 978-1-4613-7193-9 © 1999 Springer Science+Business Media New York Originally published by Kluwer Academic / Plenum Publishers, New York in 1999 Softcover reprint of the hardcover 1st edition 1999

AU rights reserved

No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written , permission ftom the Publisher ProfessorAsuncion Linares, Spanish pioneerammonitologist, has belonged to the University ofGranada since 1947, where she has been a full professor ofpalaeontology since 1961 and professor emeritus from 1987. Asuncion Linares has more than 50 years ofacademic work in Granada, including positions ofVice Rector, Director ofthe Instituto de Ciencias de la Educacion and Head ofthe Direccion de Asistencia Estudiantil at the University ofGranada. In 1960, Asuncion Linares published in Notas y Comunicaciones del Instituto Geologico y Minero de Espana and the Memoire hors-serie de la Societe Geologique de France, in memory ofProfessor Paul FaIlot, her fIrst papers on Lower ammonites gathered near Granada. At present, Professor Asuncion Linares continues 38 years ofresearch on ammonites with active work in southern Spain on mainly Middle Jurassic ammonites from the Betic Cordillera. PREFACE

Part I: Development and Evolution. S.v. Boletzky uses the reproductive system of octopods, the lower beaks of coleoids and ammonites, and the arm crown of the coleoids to analyse ontogenetic patterns and processes from an evolutionary standpoint. The author seeks to update paleomorphology to provide historical perspective for the modern biology of cephalopods. K. Uchiyama and K. Tanabe describe their hatching experiments on macromphalus in the Toba Aquarium, demonstrating the relationships between temperature and the duration of hatching periods, and establishing comparisons with Nau• tilus belauensis. E. Lefkaditou and P. Kaspiris compare analyses of morphologic features in two recent species of the genus Sepietta, revealing significant differences in club length and tentacle width, the latter expressed as a percentage ofbody weight. The monophyly of the is investigated by S.M. Klofak and collabora• tors through microscopic analyses of sculpture and internal morphology in Lower De• vonian ammonitellas (Agoniatitida) from North America and Africa. These researchers conclude that there was a non-concretionary mode of formation, which they use as evi• dence to support the monophyly of the Ammonoidea. L.A. Doguzhaeva and collaborators provide precise data concerning prismatic and nacreous layers in longiconic spirulid cephalopods from southern mid-continent using ultrastructural studies, and discuss the significance of these data for new proposals on systematics and evolution of Palaeozoic Spirulida, as well as for the clarification of problems central to evolution. The rapid diversification of monophyletic clades in restricted areas, "species flocks", is identified by M.M. Yacobucci, which recognizes the extreme plasticity in de• velopmental timing in relation to high autapomorphy:synapomorphy ratio and parallelism in Cenomanian ammonites. This author proposes that developmental plasticity rather than sensitivity is a root cause for high speciation rates in ammonites. W.L. Manger and col• laborators interpret gigantism in rich Middle ammonoid assemblages lack• ing embryonic shells. Giant specimens are interpreted as pathological, resulting from failures to attain sexual maturity in conspecific assemblages inolved in reproductive events. S.R. Fernandez-Lopez and collaborators apply biostratigraphy and palaeobiogeogra• phy to differentiate behaviour in two species of Tmetoceras colonizing epicontinental and oceanic environments in Iberia during Aalenian times. Precise biostratigraphy is employed by A. Dagys to revise the phylogeny ofTriassic Sibiritidae from Siberia and to describe the new genus Epiboreoceras. On the same basis ofbed-by-bed sampling, V.V. Mitta uses shell

vii viii Preface structure and sculpture of the inner whorls in Callovian ammonites to revise the phyletic line -Cadochamoussetia-Chamoussetia. to investigate homeomorphy and to identify the new genus Eckhardites with a probable origin in Arctocephalitinae. A.H. King provides an updated review ofVolkhovian and Kundan from Sweden, assigned to the Ellesmerocerida, , , , and ; several ofthese taxa are considered to be key forms in deciphering the evolution ofOrdovi• cian cephalopods and in providing further evidence for the systematic position of lituitid nautiloids. N. Mariotti and J.S. Pignatti revise the systematics of Xiphoteuthididae on the basis ofdiagnostic characters, taxonomic relationships and .

Part II: Form. Construction, and Function. Ultrastructural analyses are developed by S. Lenz to describe cell types in the epidermis ofthe ventral funnel and mantle ofadult vulgaris, to interpret fine structure and function, and to evaluate the similarity of this species with other molluscs. P. Ruth and collaborators investigate cytomorphologic similarities in the special cells involved in the haemocyanin synthesis in and Nauti• lus on the basis ofcomparative immunohistochemical and immunocytochemical investiga• tions. The authors identified the precise location of these special cells in branchial and midgut glands in Sepia, as well as in Nautilus where the cells are located in the midgut gland and scattered in the tissue of the ligamentum branchiale in the gill complex. H. Schmidtberg analyses functional morphology of suckers in Eledone moschata and planktic paralarvae of Octopus vulgaris by ultrastructural analysis, and establishes differ• ences in correlation with developmental types in octopod cephalopods; the author con• cludes that suckers in the taxa studied show significant differences in number and structure, as well as in the muscular and nervous systems. Ultrastructural analyses has been also applied to exceptionally preserved Conobelus (Duvaliidae) collected by L.A. Doguzhaeva and collaborators from the Valanginian ofthe Crimean Peninsula. Their study reveals differences in sublayers in the prismatic proto• conch wall, the thickness of the closing membrane, the structural differences between the primordial rostrum and the rostrum, the exact location of the replacement of retro- to pro• choanitic septal necks, and favours ultrastructural comparisons of early shells in belemni• tids and bactritoids. C. Kulicki characterizes Phylloceratina by means of aggregation patterns of needle-like crystals in the outer prismatic layer, and demonstrates that contemporaneous Lyto- and show a pseudohexagonal trilling which re• sembles nacreous plates in other molluscs. N.H. Landman and collaborators analyse empty shells ofLate Carboniferous Glaphyrites to identify typical features ofthe early on• togeny; the authors find muscle scars and ridges for proseptum attachment to be similar to those known in Mesozoic ammonoidea and propose that this could indicate similarities in their embryonic development. Investigation on the in recent and fossil cephalopods, carried out by L.A. Doguzhaeva and collaborators, reveals a similar cicatrix in the apical shell of the Lower Carboniferous and recent Nautilus, while a cup-shaped and ventrally flattened shell primordium with a marginal bilobed structure, which is typical in Rayonno• ceras, is unknown in Nautilus; these authors interpreted the observed differences as prob• able signs ofspecial modes ofembryonic shell secretion. Numerical analyses were developed to approach the description of shell features in ammonites. Shell shape and sculpture in Lower Jurassic Ammonitina (Harpoceratinae) were examined by P. Neige through the application of landmarks and geometrical techniques, which are proposed as useful tools for mapping shell features providing comparative analy• ses in ammonites. Paying attention to assumed main colonized environments, F. Oloriz and Preface ix collaborators approach the analysis of relationships between suture complexity and other shell features in Upper Jurassic ammonoids on the basis of fractal and multivariate (PCA) analyses, and evaluate the statistical significance ofthe relationships investigated. The mor• phology and sculpture ofshells in Lower Namurian ammonites from central Asia are inves• tigated by S.Y. Nikolaeva, who identifies five morpho-groups indicating closer relationships with American midcontinent rather than with Europe; the author proposes that the relationships identified were probably not influenced by lithofacies and recognizes that ammonites with low propulsive capability were dominant among the ammonoid assem• blages studied from central Asia. C. Sarti revisits data from the literature, together with material collected from the Trento Plateau (Italy), to investigate on the significance of the whorl-width:diameter ratio in order to identify dimorphic pairs in ammonites.

Part III: Taphonomy, Palaeoecology, Palaeobiogeography, Biostratigraphy, and Basin Analysis. Possible prints of the hyponome and crop-content in Lower are interpreted by H. Summesberger and collaborators in relation to rollmarks and clusters of minute brachiopods, respectively. W.L. Manger and collabora• tors interpret the unusual abundance of ammonoids with neither size selection nor evi• dence for significant post-mortem transport, ammonitellas and protoconchs, as the possible evidence of mass-mortality events following reproduction, which were recorded in both high- and low-energy deposits of Middle Carboniferous age in northern Arkansas. K. Histon uses the combined analysis of taphonomic, palaeoecologic and bathymetric data gathered from nautiloids and facies at the Rauchkofel Boden section (Carnic Alps) to interpret changing energy and oxygenation during deposition of Wenlock to Pri• moldi Cephalopod . Following methods previously developed by Hewitt and Westermann (1988), K. Histon and M. Gnoli work on a bathymetric analysis of 28 spe• cies of nautiloids revealing fluctuating environments for and cyrtonones pre• served in the " limestones" of the Upper Silurian in Sardinia; the authors conclude that deeper water conditions prevailed than previously envisaged, depths being shallower during Wenlock to Ludlow than during Primoli times. S.R. Fernandez-Lopez and collaborators use taphonomy and geographic ranges to interpret biogeographic dy• namics in Tmetoceras and separate the epicontinental species T regleyi and T flexi• costatum from the ubiquitous T scissum; on the basis of biostratigraphic interpretations, these authors identify differences in survival between eastern Pacific and western Tethys populations ofthe ubiquitous species T. scissum. New records of in the Neuquen and Austral Basins of Argentina are re• ported by P. Doyle and D. Pirrie, who discuss the significance of these records in the in• terpretation of palaeobiogeographic patterns in belemnites. E. Avram presents a re-evaluation of the record ofDeshayesites to restore its geographic distribution along the northern slope of the Tethys during the Early Aptian, and proposes the Carpathian area as the link between West-European and Caspian regions. F. Oloriz and collaborators revise ammonite biostratigraphy, providing the precise ranges ofKossmatia, Durangites and Substeuroceras in the type-area ofAlamitos in Sierra de Catorce in Mexico, show the discontinuous record ofammonite assemblages in relation to fluctuations in relative sea level, and recognize stratigraphic gaps associated with sig• nificant but laterally variable events of condensed deposition in the Alamitos area. O.K. Bogolepova and A.P. Gubanov interpret mass accumulations ofcephalopods in nar• row belts with deposition of black shales as resulting from the weakening ofoceanic gyres in outer-shelf margins during events ofupwelling affecting the East-Siberian Basin. x Preface

Lower Carboniferous biostratigraphy in Northern Russia is updated by L.F. Kusina studying at the genus level, in terms ofthe global ammonoid zones proposed by Ruzhencev and Bogolovskaya (1971); the author identifies the cosmopolitan character of a major part of the ammonoids studied, and reports differences in ranges between eastern Siberia and northeastern Europe. M.B. Aguirre-Urreta and P.F. Rawson examine the range of Valanginites in the Neuquen Basin (Argentina) and interpret differences in the range of this genus in South America and Europe in relation to differential success during short-lived migrations to peripheral regions.

Federico 0I6riz Francisco 1. Rodriguez-Tovar ACKNOWLEDGMENTS

The editors are especially indebted to the reviewers who contributed to this volume with a generous and valuable work concerning the selection and improving ofthe submit• ted papers: K. BANDEL (Geol.-Paliiont. Inst. Univ. Hamburg); R.T. BECKER (Inst. Paliiont., Humboldt-Univ., Berlin); S.v. BOLETZKY (CNRS Observ. Ocean. de Banyuls, Lab.Arago, Banyuls-sur-mer); R. BUSNARDO (Univ. Claude Bernard, Lyon); lH. CAL• LOMON (University College, London); E. CARIOU (Univ. Poitiers); lA. CHAMBER• LAIN, Jr. (Dept. Geology, Brooklyn College, NY); R. COMBEMORELL (Univ. Claude Bernard, Lyon); l COPE (Dept. Geology, Univ. Wales); G. DIETL (Staat. Mus. Naturk., Stuttgart); l-L. DOMMERGUES (Univ. Bourgogne, Dijon); S. ELMI (Univ. Claude Ber• nard, Lyon); R. ENAY (Univ. Claude Bernard, Lyon); T. ENGESER (Geol.-PaHiont. lost. Univ. Hamburg); S. FERNANDEZ-LOPEZ (Univ. Complutense, Madrid); W.M. FUR• NISH (Dept. Geology, Univ. Iowa); J. GUEX (lnst. Geol., Univ. Laussane); P. HAN• TZPERGUE (Univ. Claude Bernard, Lyon); R. HEWITT, Ph. HOEDEMAKER (Nat. Museum, Leiden); M.R. HOUSE (Dept. Geology, Univ. Hull); D.K. JACOBS (Dept. of Biology, Univ. California at Los Angeles); D. KORN (Geol.-PaHiont. Inst., Tiibingen); l KULLMANN (Geol.-PaHiont. Inst., Tiibingen); N.H. LANDMAN (Dept. of Invertebrates, Amer. Mus. Nat. Hist., NY); D. MARCHAND (Univ. Bourgogne, Dijon); l MUTTER• LOSE (Inst. Geol. Paliiont., Univ. Hannover); T. OKAMOTO (Dept. Earth Sci., Ehime Univ.); RAUP (Dept. Geophysical Sci., Univ. Chicago); G.R. STEVENS (New Zealand Geol. Survey); P.F. RAWSON (Dept. Geol. Sci., Univ. College London); A. RICCARDI (Univ. Nac. La Plata, Buenos Aires); A. SALVADOR (Dept. Geol. Sci., Univ. Texas at Austin); A. SEILACHER (Geol.-PaHiont. Inst., Tiibingen); K. TANABE (Geol. Inst. Univ. Tokyo); J. THIERRY (Univ. Bourgogne, Dijon); E.T. TOZER (Geological Survey ofCan• ada); G.E.G. WESTERMANN (McMaster Univ., Hamilton); and A. ZEISS (Inst. Palaont., Friedrich-Alexander-Univ., Erlangen).

xi CONTENTS

Part I: Development and Evolution

I. Cephalopod Development and Evolution: Biological Insight into Ontogenesis as a Guide to Paleomorphology 3 S. v. Boletzky

2. Hatching ofNautilus macromphalus in the Toba Aquarium, Japan...... 13 Kimio Uchiyama and Kazushige Tanabe

3. Comparative Analysis ofSome Morphometric Characters in Two Sepiolids: Sepietta neglecta (Naef, 1916) and Sepietta oweniana (Orbigny, 1840) ... -17 E. Lefkaditou and P. Kaspiris

4. Embryonic Development ofPrimitive Ammonoids and the Monophyly ofthe Ammonoidea ...... 23 S. M. Klofak, N. H. Landman, and R. H. Mapes

5. A Late Carboniferous Spirulid Coleoid from the Southern Mid-Continent (USA): Shell Wall Ultrastructure and Evolutionary Implications 47 Larisa A. Doguzhaeva, Royal H. Mapes, and Harry Mutvei

6. Plasticity ofDevelopmental Timing as the Underlying Cause ofHigh Speciation Rates in Ammonoids: An Example from the Cenomanian Western Interior Seaway ofNorth America...... 59 Margaret M. Yacobucci

7. Pathologic Gigantism in Middle Carboniferous Cephalopods, Southern Midcontinent, United States 77 Walter L. Manger, Lisa K. Meeks, and Daniel A. Stephen

8. Aalenian Tmetoceras (Ammonoidea) from Iberia: Taxonomy, Habitats, and Evolution ...... 91 S. R. Fernandez-Lopez, M. H. Henriques, A. Linares, 1. Sandoval, and M. S. Ureta

xiii xiv Contents

9. Evolution ofthe Family Sibiritidae and Detailed Biostratigraphy ofthe Siberian Upper Olenekian () . 109 Algirdas Dagys

10. The Genus Cadochamoussetia in the Phylogeny ofthe Jurassic (Ammonoidea) . 125 Vasilii V. Mitta

II. A Review ofVolkhovian and Kundan (-Llanvirn) Nautiloids from Sweden 137 Andrew H. King

12. The Xiphoteuthididae Bather, 1892 (, ): An Outline Classification . 161 Nino Mariotti and Johannes S. Pignatti

Part II: Form, Construction, and Function

13. Ultrastructural Studies on the Epidermis ofAdult Octopus vulgaris Cuvier, 1797 173 Sabine Lenz

14. Comparative Immunohistochemicaland Immunocytochemical Investigations on the Location ofHaemocyanin Synthesis in Dibranchiate and Tetrabranchiate Cephalopods (Sepia and Nautilus) 189 P. Ruth, R. Schimmelpfennig, and R. Schipp

15. Ultrastructural Studies ofthe Suckers ofNewly Hatched Eledone moschata and Octopus vulgaris (; Cephalopoda) ...... 203 Henrike Schmidtberg

16. Conch Ultrastructure and Septal Neck Ontogeny ofthe Belemnite Conobelus (Duvaliidae) from the Valanginian ofthe Crimea (Black Sea) 223 Larisa A. Doguzhaeva, Harry Mutvei, George K. Kabanov, and Desmond T. Donovan

17. Aggregation ofAragonite in the Outer Prismatic Layer ofMesozoic Ammonoids 233 Cyprian Kulicki

18. Internal Features ofthe Embryonic Shells ofLate Carboniferous Goniatitina 243 Neil H. Landman, Royal H. Mapes, and Kazushige Tanabe

19. Early Ontogeny ofthe Siphuncle and Shell in the Early Carboniferous Rayonnoceras (Actinocerida) from Arkansas, USA 255 Larisa A. Doguzhaeva, Harry Mutvei, and Royal H. Mapes

20. The Use of Landmarks to Describe Ammonite Shape: Examples from the Harpoceratinae 263 Pascal Neige Contents xv

21. Recent Advances in Morphometric Approaches to Covariation ofShell Features and the Complexity ofSuture Lines in Late Jurassic Ammonites, with Reference to the Major Environments Colonized 273 Federico Oloriz, Paul Palmqvist, and Juan A. Perez-Claros

22. Morphological Diversity ofAmmonoids from the Lower Namurian ofCentral Asia...... 295 Svetlana V. Nikolaeva

23. Whorl Width in the Body Chamber ofAmmonites as a Sign ofDimorphism 315 Carlo Sarti

Part III: Taphonomy, Palaeoecology, Palaeobiogeography, Biostratigraphy, and Basin Analysis

24. Rollmarks ofSoft Parts and a Possible Crop Content ofLate Ammonites from the Slovenian Karst 335 Herbet Summesberger, Bogdan Jurkovsek, and Tea Kolar-Jurkovsek

25. Possible Cephalopod Reproductive Mass Mortality Reflected by Middle Carboniferous Assemblages, Arkansas, Southern United States 345 Walter L. Manger, Daniel A. Stephen, and Lisa K. Meeks

26. Silurian Cephalopod Facies in the Carnic Alps (Rauchkofel Boden Section, Austria): Taphonomy ofthe Fauna...... 365 Kathleen Histon

27. Nautiloid Paleobathymetry from the Silurian 'Orthoceras Limestone' Facies of SW Sardinia, Italy ...... 381 Kathleen Histon and Maurizio Gnoli

28. Aalenian Tmetoceras (Ammonoidea) from Iberia: Taphonomy and Palaeobiogeography 395 S. R. Fernandez-Lopez, M. H. Henriques, A. Linares, 1. Sandoval, and M. S. Ureta

29. Belemnite Distribution Patterns: Implications ofNew Data from Argentina 419 Peter Doyle and Duncan Pirrie

30. The Deshayesites Kazansky, 1914 (Ammonoidea) Representatives in Romania, a Link between the West-European and Caspian Assemblages ofthis Genus.. 437 Emil Avram

31. Ammonite Biostratigraphy and Correlations in the Upper Jurassic-Lowermost Cretaceous La Caja Formation ofNorth-Central Mexico (Sierra de Catorce, San Luis Potosi) 463 F. Oloriz, A. B. Villasenor, C. Gonzalez-Arreola, and G. E. G. Westermann xvi Contents

32. Silurian Cephalopod Limestone Biofacies from Eastern Siberia: Fauna, Age, and Environments 493 Olga K. Bogolepova and Alexander P. Gubanov

33. Biostratigraphic Distribution ofLower Carboniferous Ammonoids in Northern Russia 505 Lydia F. Kusina

34. Stratigraphic Position of Valanginites, Lissonia, and Acantholissonia in the Lower Valanginian (Lower Cretaceous) Ammonite Sequence ofthe Neuquen Basin, Argentina . 521 Maria B. Aguirre-Urreta and Peter F. Rawson

Index 531