Constructional Morphology and Evolution

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2 N. Schmidt-Kittler' K. Vogel (Eds.) Constructional Morphology and Evolution With 147 Figures Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Barcelona Budapest

3 Prof. Dr. NORBERT SCHMIDT -KITTLER Institut fur Geowissenschaften Johannes Gutenberg-UniversiUit Saarstr Mainz, FRG Prof. Dr. KLAUS VOGEL Geologisch-Palaontologisches Institut J. W. Goethe-Universitat Senckenberganlage Frankfurt a. M., FRG Cover illustration (drawn by H. Schafer): The cover illustration symbolizes a basic principle of constructional morphology: a body surrounded by a flexible envelope and tightly filled with fluid or viscid material automatically assumes a spherical shape. This is true for protists as well as for multicellular organisms. Every deviation must be actively generated by a bracing system. Ontogeny and phylogeny are governed by this principle. This holds also for skeleton bearing organisms such as trilobites. Skeletal growth requires preformed hydraulic bodies. In trilobites as in other arthropods this reciprocal effect of hydraulics and bracing systems also maintains the shape during molting processes. ISBN-13: e-isbn-13: DOl: / This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks. Duplication of this publication or parts thereof is only permitted under the provisions of the German Copyright Law of September 9, 1965, in its current version, and a copyright fee must always be paid. Violations fall under the prosecution act of the German Copyright Law. Springer-Verlag Berlin Heidelberg 1991 Softcover repint of the hardcover 1 st edition 1991 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use Printed on acid-free paper

4 Preface This book contains the results of an international symposium held in Bad Homburg, FRO, from the 17th to the 21st April 1990 under the title "Constructional Morphology and Biomechanics: Concepts and Implications". 1\venty-five biologists and paleontologists from different countries working in this field were invited to present review papers and case studies on the main fields of constructional morphology. The list of participants is practically identical with the list of contributors given on page XI with the exception that in co-authored works only the first author attended. The contributions of the participants were submitted before the meeting took place so that the symposium could mainly be dedicated to discussion. All contributions were reconsidered by their authors after the symposium in response to discussion remarks and after having been reviewed by one or two other participants of the meeting. The articles are divided into four main topic fields: Papers dealing with conceptual and methodical questions of constructional morphology form the first part. It is followed by a group of papers focussing on the operation of bioconstructions and behavioural control within the environment. Morphogenetic aspects of bioconstructions and organic patterns are the subject of the third part. The last part focusses on the evolution of bioconstructions and body plans. Some of the contributions, due to their more general approach or the combination of treated subjects, could have been assigned to more than one topic field. Thus, their arrangement in one part of the book must seem somewhat arbitrary, however, a decision had to be made. The symposium was financed by the Werner Reimers-Stiftung in Bad Homburg and the Siftung Volkswagenwerk in Hannover, both of which are thanked for their support. We are also grateful to the President of the University of Mainz for his financial contributions to the organization of the symposium. The very pleasant and relaxing atmosphere of the facilities and grounds of the Reimers-Stiftung in Bad Homburg was an ideal meeting place for scientific encounters of a small working group. March 1991 Norbert Schmidt-Kittler, Mainz Klaus Vogel, Frankfurt

5 Contents Introduction: A New Look at Morphology N. Schmidt-Kittler and K. Vogel.... Part I Concepts of Constructional Morphology Explanations in Konstruktionsmorphologie and Evolutionary Morphology W.J. Bock... 9 Functional Aspects of Morphology W. Nachtigall An Outline of Morpho-Constructional Analysis N. Schmidt-Kittler Concepts of Constructional Morphology K. Vogel Part II Bioconstruction: Operational Interconnections, Behavior, Environment Dynamic Similarity in the Analysis of Animal Movement R. McN. Alexander Cytomechanics and Biochemistry J. Bereiter-Hahn Constructional Principles and the Quasi-Experimental Approach to Organisms W.E Gutmann Biomechanics in the Enamel of Mammalian Teeth W. v. Koenigswald and H.-V. Pfretzschner The Functional Morphology of Gastropod Apertures C. R. C. Paul Behavior plus "Pathology" - the Origin of Adaptations? D. S. Peters The Function of Labyrinthodont Teeth: Big Teeth in Small Jaws H. Preuschoft, W.-E. Reif, C. Loitsch and E. Tepe

6 VIII Contents Complexity and a Coupled System: Flight, Echolocation and Evolution in Bats J. M. V. Rayner Biomechanical Aspects of the Wing Joints in Flies, Especially in Calliphora erythrocephala A. Wisser and W Nachtigall Part III Bioconstruction: Morphogenesis Ontogenetic Changes Reflected in the Morphology of the Molluscan Shell K. Bandel A Functional Approach to the Development of the Head of Teleosts: Implications on Constructional Morphology and Constraints K. F. Liem Self-Organizing Mechanisms in Morphogenesis and Evolution A. Seilacher Genetic and Molecular Analysis of Pattern Formation Processes in Drosophila D. Tautz Design Elements Employed in the Construction of Animal Skeletons R. D. K. Thomas and W-E. Reif A Theoretical Morphological Approach to Tooth Replacement in Lower Vertebrates D. B. Weishampel Part IV Bioconstruction: Evolution Evolution of Biological Constructions: Concessions, Limitations, and Pathways P. Dullemeijer Constructional and Ecological Prerequisites for the Evolution of Homeothermy H.-R. Duncker The Mechanical Constraints in Mollusc Constructions the Function of the Shell, the Musculature, and the Connective Tissue K. Edlinger Morphogenetic Versus Environmental Cues for Adaptive Radiations P. W Skelton

7 Contents IX The Sequence of Body Plans and Locomotory Systems During the Precambrian-Cambrian Transition J. W. Valentine Subject Index 399 Index of Taxonomic Terms

8 List of Contributors You will find the addresses at the beginning of the respective contribution Alexander, R.McN. 71 Bandel, K. 211 Bereiter-Hahn, J. 81 Bock, W.J. 9 Dullemeijer, P. 313 Duncker, H.-R. 331 Edlinger, K. 359 Gutmann, W. F. 91 von Koenigswald, W. 113 Liem, K. F. 231 Loitsch, C. 151 Nachtigall, W. 31, 193 Paul, C. R. C. 127 Peters, D. S. 141 Pfretzschner, H.-V. 113 Preuschoft, H. 151 Rayner, J. M. V. 173 Reif, W.-E. 151, 283 Schmidt-Kittler, N. 1, 41 Seilacher, A. 251 Skelton, P. W. 375 Tautz, D. 273 Tepe, E. 151 Thomas, R. D. K. 283 Valentine, J. W. 389 Vogel, K. 1, 55 Weishampel, D. B. 295 Wisser, A. 193