Adolf Seilacher Trace Fossil Analysis Adolf Seilacher

Trace Fossil Analysis

With 75 Plates and 43 Photos Autor Prof. Dr. Adolf Seilacher Yale Geology Dept. P.O. Box 208109 New Haven, CT 06520-8109 USA and Engelfriedshalde 25 72076 Tübingen Germany E-mail: [email protected]

Library of Congress Control Number: 2006935138

ISBN-13 978-3-540-47225-4 Springer Berlin Heidelberg New York

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Printed on acid-free paper 32/2132/CA – 5 4 3 2 1 0 Preface

This is a course book – meaning that it intends to confer not knowledge, but skill. The need for this skill becomes obvious if we look at the changing role of trace fossils during the last decades. From objects that were treated in standard paleontology textbooks, at best, under “Miscellanea”, together with problematica, coprolites and pseudofossils useless as index fossils, they have become subject of a special field, paleoichnology. A journal (ICHNOS), Ichnological Newsletters and regular workshops have been established, symposia are held, and the literature has increased exponen- tially. This success stems mainly from the intimate connection of ichnology with sedimentology and the importance that both fields have for paleoenvironmental and ba- sin analysis, which becomes more and more important in petroleum exploration. This useful connection, however, also had its price. In the hands of biogeologists, trace fossils easily loose their significance as unique biological documents. They are commonly treated summarily as “bioturbation”, a term that was originally meant to describe the biogenic destruction of primary depositional structures and the lack of distinctive trace morphologies. In a more quantitative form, “ichnofabrics”, this approach has had considerable success. On the taxonomic side, paleoichnology is all too often considered as a field, in which there is no limit to coining new names and taxa without affiliations. If some- body would take the effort today to revise Walter Häntzschel’s trace fossil volume of the “Treatise on Invertebrate Paleontology”, it would probably double in size. Still a large proportion of the ichnogenera described in the meantime would probably fall under “unrecognized” or “synonyms”. The situation being as it is, this book bypasses the necessary job of taxonomic revision (the divergence between lumpers and splitters is extremely wide in this field). Rather it concentrates on the more distinctive and representative ichnogenera. It is also focussed on structures left by invertebrate animals in soft sediments, treating vertebrate tracks only marginally and leaving out hard substrate borings, bite marks on body fossils, eggs, nests, and coprolites altogether. Not that such documents would not merit attention; but by their very nature they require a different approach and should better be treated, together with the respective body fossils, by systematic paleontologists. A word is also necessary about how this book originated and how it should be used. It grew out of courses I gave in Tübingen and many other places with the aid of representative specimens and plaster casts. In the printed version, the plates remain the core, to which the text is added in the form of extended captions. It remains to be seen, how far the style of this course can be detached from my own personal interac- tion and how far colleagues with different views and experiences are willing to adopt it – if only to develop alternative and possibly more adequate ideas. In no case, however, should it be done without real material, because it is the inquisitive observation that should be trained. In my own experience, drawing specimens with an old-fash- ioned camera lucida is still the most adequate method. If a form appears to be too complicated, more often than not it will become understandable after our brain has absorbed it through the pencil. Many of my drawings are new, others repeated from VI Preface

earlier publications; but all of them have been arranged in such a way that they make a story and wait for the addition of your own colorful scribblings. This also means that you should devote more than a glimpse to these illustrations. After all, early illiterate periods conveyed whole world views through pictures. For the student that wants to delve further into a subject, relevant papers are arranged by plates and annotated. A glossary explains the terminology. There follow the apologies: for not covering all ichnogenera (which can be found in the “Treatise”); for covering too much in a too condensed style (objection of my wife); and for unrealistically assuming that the time of fifteen two-hour courses could be spared for trace fossils in the curriculum of any school. But again, it is not the knowledge of forms and names that we aim at, but a method of morphological think- ing in terms of processes that could easily be transferred to any other subject matter. So let us follow the motto of my Tübingen University: ATTEMPTO (Let us try!). Acknowledgements

I collected my first trace fossils 68 years ago. Listing acknowledgements over such a long timespan amounts to writing one’s own biography. My tutors in highschool times were a local physician , Dr. R. Stierlin, and a devoted forester, Dr. h.c. Otto Linck. Back at Tübingen University for the first post-war semester, the famous vertebrate paleon- tologist Prof. F. von Huene taught me to use the camera lucida, which has since re- mained my most important aid. Nevertheless it was the broader-minded Prof. Otto H. Schindewolf, who encouraged me to do the doctoral thesis on Mesozoic trace fossils. This brought me in contact with the Senckenberg Institute in Wilhelmshaven, where I was introduced to actuo-paleontology by Prof. Wilhelm Schäfer, who also influenced my drafting style, and with Prof. Walter Häntzschel, who then compiled the Treatise volume on “Trace Fossils and Problematica”. Equally important was a first trip to Italy, where the rich university collections in Pisa and Florence made me familiar with the strange flysch ichnocoenoses and with the new ideas of Prof. C. I. Migliorini about sand being imported into deepsea basins. Later, the recogni- tion of the Nereites ichnofacies lent strong support to his and P. H. Kuenen’s turbidite theory, as well as to the new paradigm of plate tectonics. Right after promotion (1951), I had the priviledge to join my teacher on his expe- dition to the Salt Range of Pakistan. Although our original target had been the end- Permian mass extinction, this trip introduced me to Cambrian shallow-marine ichnocoenoses, in which the various activities of trilobites play a dominant role. Later field work in Spain, in which the advise of Prof. F. Lotze has been crucial, led to the scheme of Cruziana ichnostratigraphy. During the following years it was successfully applied to the dating of otherwise non-fossiliferous sandstones in the deserts of North Africa, where E. Klitzsch and his team were of invaluable help. It also stood the test in a memorable excursion to the Tassili arranged by the Algerian Geological Survey in order to study the effects and the dating of the newly discovered end-Ordovician glaciation. Ichnostratigraphy was further refined in the symposia of the National Oil Company of Libya organized by M. Salem, trips to Jordan arranged by F. Bender, and to Saudi Arabia with M. Senalp. On the American side, the interest of oil companies was more in facies relation- ships of trace fossils. Consulting for Jersey Research Company by the initiative of J. Campbell gave me a chance to study a broad range of ichnocoenoses through the south-western part of the country. The idea to compound this book came with the invitation by X. T. WU to hold a course in Jao Zuo (Henan, China) with more than hundred participants. It gradually matured during similar courses throughout the world; but only when presenting it with Derek Briggs at Yale University (where I am teaching since retirement from Tübingen University ), he gave me the ultimate kick to finish the manuscript. Several people have been involved in its completion. Our son Peter Seilacher cleaned the illustrations with the computer, Gabriela Mangano, Luis Buatois (University of Saskatchewan) and Andrew Rindsberg (University of Georgia) helped in accumulat- ing and annotating the list of references. Above all, Edith Seilacher-Drexler, my dear VIII Acknowledgements

wife and unpaid partner for half a century, did all the word processing involved in updating and re-arranging the text. The late Roland Goldring, Gabriela Mangano, Luis Buatois, and Andrew Rinds- berg were of tremendous help in compiling and annotating the relevant references. At various points, Wolfgang Gerber, Hans Luginsland, Werner Wetzel, Eden Volohonsky (Tübingen), Bill Sacco (New Haven) and Jens Rydell (Göteborg) supplied photographs. Hans Luginsland also produced casts in the field and in his small labo- ratory. Last not least I am obliged to Springer-Verlag, whose representative Wolfgang Engel kept encouraging me throughout the years to finish this book but did not live to see it published. During the fruitful collaboration with Armin Stasch (Bayreuth) I learned about the crucial function of a scientific book designer in producing a book. My deep gratitude goes to these people, not to forget all the inquisitive students in the courses. May their future generations share my contention that it is a wonderful thing to work as a paleodetective! General References

Textbooks

Richter R (1925) Flachseebeobachtungen zur Paläontologie und Geologie. III–VI. Senckenberg 8(3/4): 200–224 Abel O (1935) Vorzeitliche Lebensspuren. Fischer, Jena (A classic integrative text that non-German readers will nevertheless enthuse on) Bromley RG (1996) Trace fossils: biology, taphonomy and applications, 2nd edn. Chapman & Hall, London, 361 p (A very readable text with good appreciation of modern traces and the palaeobiology of ancient ones. Rather thin sedimentologically and does not discuss hardgrounds. Includes a glossary) Ekdale AA, Bromley RG, Pemberton SG (1984) Ichnology – the use of trace fossils in sedimentology and stratigraphy. Society of Economic Paleontologists & Mineralogists (SEPM), Short Course 15, 317 p Ekdale AA, Pemberton SG, Bromley RG (1984) Trace fossils. Society of Economic Paleontologists & Mineralogists (SEPM). Short Course 15, 317 p (A useful but dated text for sedimentological applications) Frey RW (ed) (1975) The study of trace fossils. Springer-Verlag, Heidelberg, 562 p (A rather dated text) Lockley MG (1991) Tracking dinosaurs: A new look at an ancient world. Cambridge University Press, Cambridge, 238 p Maples CG, West RR (eds) (1992) Trace fossils: Short courses in paleontology, 5. Paleontological Society, Cincinnati Schäfer W (1972) Ecology and palaeoecology of marine environments. Oliver and Boyd, Edinburgh, 568 p (This text, superbly translated from the German edition is in the truly scientific spirit of careful observation and interpretation. It concerns observations made predominately in the intertidal zone of the southern North Sea)

Source Books

Crimes TP, Harper JC (eds) (1970) Trace fossils. Geol J, Special Issue, 3, Seel House Press, Liverpool (A collection of 35 mostly systematic papers) Crimes TP, Harper JC (eds) (1977) Trace fossils 2. Proceedings of an International Sympopsium held at Sydney, Australia. Geol J, Special Issue, 9, Seel House Press, Liverpool (A collection of 15 mostly systematic papers) Donovan SK (ed) (1994) The palaeobiology of trace fossils. The John Hopkins University Press, Bal- timore, 308 p (A collection of 11 papers on some of the aspects dealt with here, and other objects such as plant roots, coprolites and vertebrate eggs) Gillette DD, Lockley MG (eds) (1989) Dinosaur tracks and traces. Cambridge University Press, Cam- bridge, 454 p (50 papers on all aspects) Häntzschel W (1965) Vestigia Invertebratorum et Problematica. Fossilium Catalogus 1: Animalia 108, W. Junk, s’Gravenhage, 142 p (Most complete annoteted bibliography of the time) Häntzschel W (1975) Trace fossils and problematica. In: Teichert C (ed) Treatise on invertebrate paleontology, Part W, Supplement 1. Geological Society of America and Univerity of Kansas, W1– W269 (The A–Z of trace fossils and related structures) Kuhn O (1963) Ichnia tetrapodorum. Fossilium Catalogue, 1, Animalia 101, W. Junk, s’Gravenhage, 176 p (With 12 plates of line figures) Lessertisseur J (1955) Traces fossiles d’activitié animale et leur significance paléobiologique. Soc Geol Fr Mem New Ser 74:1–150 (General review of invertebrate trace fossils) Miller III W (ed) (2003) New interpretations of complex trace fossils. Palaeogeog Palaeoclim Palaeoecol (Special issue) 192, 343 p (18 papers by paleontologists and biologists concerned with the functional and fabricational context of invertebrate burrow systems. A most stimulating approach as opposed to the one of ichnofabrics) X General References

Miller MF, Ekdale A, Picard MD (eds) (1984) Trace fossils and paleoenvironments: Marine carbon- ate, mariginal marine terrigenous and continental terrigenous. J Paleontol 58(2):283–597 (22 papers on invertebrate trace fossils) Noda H (1982) Check list and bibliography of trace fossils and related forms in Japan (1889–1980) and neighbourhood (1928–1980). Introduction to study of trace fossils, part 2, pp 1–80, Ibaraki, Japan Seilacher A (1997) Fossil art. An exhibition of the Geologisches Institut Tübingen University. The Royal Tyrell Museum of Palaeontology, Drumheller, Alberta, Canada, 64 p (Large casts of trace fossils and other sedimentary structures, arranged for their appeal)

General Papers

Häntzschel W (1955) Rezente und fossile Lebensspuren, ihre Deutung und geologische Auswertung. Experientia 11(10):373–382 Richter R (1922) Flachseebeobachtungen zur Paläontologie und Geologie. III–VI. Senckenberg 4(5): 103–141 (Actuopaleontology) Richter R (1924) Flachseebeobachtungen zur Paläontologie und Geologie. III–VI. Senckenberg 6(3/4): 119–164 Contents

Chapter I · Vertebrate Tracks ...... 1

Plate 1 Chirotherium: The Sherlock Holmes Approach ...... 6 Plate 2 Undertracks in Wet Sands ...... 8 Plate 3 Tambach Ichnotope ...... 10 Plate 4 Coconino-Type Ichnotopes ...... 12 Plate 5 Fish Trails ...... 14

Chapter II · Arthropod Trackways ...... 17

Plate 6 Limulid Tracks (Kouphichnium) ...... 20 Plate 7 Other Arthropod Trackways ...... 22 Plate 8 Trilobite Tracks ...... 24 Plate 9 Adventures of an Early Cambrian Trilobite ...... 26 Plate 10 Potsdam Sandstone Trackways ...... 28

Chapter III · Trilobite Burrows ...... 31

Plate 11 Trilobite Biology and Cruziana Authorship ...... 34 Plate 12 Trilobite Fingerprints ...... 36 Plate 13 Cruziana Modifications ...... 38 Plate 14 Cruziana semiplicata ...... 40 Plate 15 Burrowing Behavior of Silurian Trilobites ...... 42

Chapter IV · Arthropod Tunnel Systems ...... 45

Plate 16 Crab and Shrimp Burrows ...... 50 Plate 17 Other Arthropod Tunnels and Nests ...... 52 Plate 18 Ophiomorphids ...... 54 Plate 19 Rhizocoralliids ...... 56 Plate 20 Rhizocoralliid Modifications ...... 58

Chapter V · Resting Traces ...... 61

Plate 21 Burrowing Techniques ...... 64 Plate 22 Undertrace Experiments ...... 66 Plate 23 Bilateral Resting Traces ...... 68 Plate 24 Asterozoan Resting Traces ...... 70 Plate 25 Coelenterate Resting Burrows ...... 72 XII Contents

Chapter VI · Burrows of Short Bulldozers ...... 75

Plate 26 Echinoid Burrows ...... 78 Plate 27 Molluscan Bulldozers ...... 80 Plate 28 Paleozoic Psammichnitids ...... 82 Plate 29 Psammichnitid Behavioral Evolution ...... 84 Plate 30 Bi- and Tripartite Backfill Bodies ...... 86

Chapter VII · Burrows of Wormlike Bulldozers ...... 89

Plate 31 Trace Fossil Classification ...... 92 Plate 32 Unbranched Burrows ...... 94 Plate 33 Spiral Burrows ...... 96 Plate 34 Nereitids ...... 98 Plate 35 Gyrochortids ...... 100

Chapter VIII · Burrows of Stripminers ...... 103

Plate 36 Modern U-Tubes ...... 106 Plate 37 Backfill Structures ...... 108 Plate 38 Post-Paleozoic Zoophycos ...... 110 Plate 39 Daedaloid Burrows ...... 112 Plate 40 Lophocteniids ...... 114

Chapter IX · Arthrophycid Burrows ...... 117

Plate 41 Teichichnid Burrows ...... 120 Plate 42 Arthrophycids I ...... 122 Plate 43 Arthrophycids II ...... 124 Plate 44 Arthrophycids III ...... 126 Plate 45 Arthrophycids IV ...... 128

Chapter X · Probers ...... 131

Plate 46 Asterosomids ...... 134 Plate 47 Medusiform Burrows (Gyrophyllitids) ...... 136 Plate 48 Various Fucoids ...... 138 Plate 49 Undermat Miners ...... 140 Plate 50 Chondritids ...... 142

Chapter XI · Deepsea Farmers ...... 145

Plate 51 Meandering ...... 148 Plate 52 Graphoglyptids I ...... 150 Plate 53 Graphoglyptids II ...... 152 Plate 54 Graphoglyptids III ...... 154 Plate 55 Paleodictyon ...... 156

Chapter XII · Pseudo-Traces ...... 159

Plate 56 Trace-like Body Fossils: Xenophyophoria ...... 162 Plate 57 Tool Marks ...... 164 Plate 58 Synsedimentary Structures ...... 166 Plate 59 Diagenetic Structures ...... 168 Plate 60 Tectograms ...... 170 Contents XIII

Chapter XIII · Earliest Trace Fossils ...... 173

Plate 61 Pre-Ediacaran Dubiostructures ...... 176 Plate 62 Ediacaran Sole Features ...... 178 Plate 63 Traces of Early Molluscs ...... 180 Plate 64 Treptichnus pedum ...... 182 Plate 65 The Cambrian Revolution ...... 184

Chapter XIV · Cruziana Stratigraphy ...... 187

Plate 66 Stratigraphic and Paleogeographic Distribution ...... 190 Plate 67 Cambrian Trilobite Burrows ...... 192 Plate 68 Ordovician Trilobite Burrows ...... 194 Plate 69 Silurian to Carboniferous Trilobite Burrows ...... 196 Plate 70 Trans-Gondwanan Seaway ...... 198

Chapter XV · Ichnofacies ...... 201

Plate 71 Global Ichnofacies ...... 204 Plate 72 Post-turbidite Ichnocoenoses ...... 206 Plate 73 Frozen Tiering and Telescoping in Shallow-Marine Settings ...... 208 Plate 74 Interactions between Trace Fossils ...... 210 Plate 75 Solnhofen Mortichnia ...... 212

Glossary ...... 215

Index ...... 217