RHAETIAN ARTICULATE BRACHIOPODS of EUROPE by DAVID AMORY BAPTY PEARSON a Thesis Submitted for the Degree of Doctor of Philosophy

RHAETIAN ARTICULATE BRACHIOPODS OF EUROPE BY DAVID AMORY BAPTY PEARSON A thesis submitted for the degree of Doctor of Philosophy of the University of London. Geology Department, Imperial College of Science and Technology. January, 1967. 2 ABSTRACT With the exception of the extremely rare thecideids and thecospirids„ all the articulate brachiopods from the Rhaetian of the Western Alps of France and Switzerland, the Northern Alps of Austria and Bavaria, the Lombardy Alps of northern Italy, the Carpathians of Slovakia, the Tatra of Poland and the Persani Mts. of northern Rumania are revised in this thesis♦ The work has been based on collections made by the writer from the whole of this area with the exception of the Western Alps. From eight superfamilies, thirty four species are described. Their relationship to those from the Norian of Slovakia and eastern Bulgaria is discussed, also using collections made by the writer, and the recently revised Norian fauna of the southern Soviet Union is considered throughout, New taxa comprise seven species, four genera and one subfamily. The composition and status of all families, subfamilies and genera is discussed in detail. In particular the content of the rhynchonellid subfamilies, Cirpinae, Rhynchonellininae and Tetrarhynchiinae is considered at length and several subfamilial divisions are suggested. Serial transverse grinding has been used to study the internal structure of all but the rarest species and photomicrographs accompany detailed drawings made from acetate peels. The ecology of many of the species is discussed and the adaptation of Austrirhynchia cornigera to orientation in currents is proposed. As a preliminary to the palaeontological study, the stratigraphy and status of the Rhaetian stage itself is discussed in detail. It is considered that the lower boundary may be usefully lowered to coincide with the disappearance of Monotis. 3 CONTENTS ABSTRACT 2 ACKNOWLEDGMENTS 5 INTRODUCTION Previous Research 7 Object of Study 9 Field Programme 10 DEVELOPMENT AND DISTRIBUTION OF THE MARINE RHAETIAN IN EUROPE 11 HISTORY AND PROBLEMS OF RHAETIAN STRATIGRAPEY 26 GENERAL PALAEONTOLOGY Ecology 1 (Brachiopod communities) 40 Ecology 11 (Adaptation of Austrirhynchia cornigera 44 Geographical Distribution 50 Stratigraphical Relationship 51 SYSTEMATIC PALAEONTOLOGY Notes 53 Laboratory Techniques 55 Terminology 56 1 REYNCTIONELLIDA Addendum 66 Rhynchonellacea Fis s irhynchia 68 Piarorhynchia 86 Gen.nov.A 87 Austrirhynchia 98 Calcirhynchia 108 ?Caucasorhynchia 126 Vindobonella 127 Osmarella 138 Caraxezzia 164 Other Rhynchonellida reported from the Rhaetian 171 4 2 SPIRIFERIDA Spiriferinacea Sinucosta 175 Laballa 183 Zugmayerella 187 Retziacea Neoretzia 206 Athyridacea Oxycolpella 211 Reticulariacea Mentzelia 220 Mentzeliopsis 225 TEREBRATULIDA Erratum 228 Zeilleriacea Zeilleria 229 Austrilleria 254 Terebratulacea Triadithyris 266 Dielasmatncea Rhaetina 277 Other Terebratulida from the Rhaetian 297 REFERENCES 298 PLATES 1-19 318 5 ACKNOWLEDIENTS The opportunity is welcomed to thank the very many people who have made this thesis possible, especially those members of Universities and Institutes in several countries who have given their time to act as hosts and to guide the writer in the field: Dr. Milos SIBLIK of the Geological Institute in Prague; Dr. Andrej RADWANSKI and Dr. George WAZEK of the University of Warsaw; Dr. Miltcho GANEV, Dr. Stefan STEPANOV, Dr. Platon TCHUNIACRENKO, Dr. Rossica KALVACHEVA. and the late Dr.Juli STEFANOV of the Geological Institute in Sofia; Dr. George M./IA-TEA, Dr. Valentin GEORGESCU and Dr. Josef a BORDEA of the Geological Institute in Bucharest; Dr. Maurizio GAETIANI of the Palaeontological Institute in Milan; Dr. Ulrich SCEWIEDRZIK of the University of Innsbruck; and Herr Ernst KATZER of Waldegg, Piesting Tal, Austria. Thanks are also due to Mr.E.OWEN of the B.M.N.E.; Professor 0.KUHN and Dr.F.STEININGER of the Pal.Inst.Univ.Viennn; Professor E.KUPPER, Professor R.SIEBER and Dr.B.PLoCEINGER of the Geol.Hundes.Vienna; Professor E.ZAPFE and Dr.H.KOLLMAN of the Nat.hist.Mus.Vienna; Professor DEEM, Dr.W.BARTHEL and Dr. ZOBELF,IN of the Bayer.St.S1g.Munich; Dr.O.GANSS of the Bayer.Geol. Landesamt, Munich; Dr.F.FABRICIUS of the TechnLche Hochschule, Munich; Dr.A.HOLZL of Hausham; Dr.M.MEEL and Mr.J.PEVNY of the Geological Institute, Bratislava; Professor A.IINDRUSOV and Mthe.Dr.W.ANDRUSOVA of the Czechoslova - kinn Academy of Sciences, Bratislava; Dr.G.VIGH of the Geological Institute, Budapest; the late Dr.G.RIIIL-N6NU and Dr.D.PATRULIUS of the Geological Institute, Bucharest; Professor W.GOETEL of the Institute of Mines, Cracow; Dr.A.D.LGIS of the Geological Institite, Novosibirsk; Dr.T.TOZER of the Geological Survey of Canada; and to Dr.A.FISCi-iER of Princeton University.. The library facilities afforded by the B.M.N.H and the Geological Society of London are also gratefully acknowledged. The whiter would also like to express his appreciation of the generous hospitality afforded to him in the field, particularly by Familie KATZLR of Waldegg, Austria; Famine METNSTER of Seegatterl, Bavaria; and Famine RUS of Satul Saud, Rumania. 6 Invaluable discussion with former colleagues, DriPaul COPPER and Dr./arm CHILDS is also gratefully ackno,11-'.ged, as is the assistance given by Mr.Harry DOUST and Miss Mary PUGH with the introductory diagrams, the advice of M±.J.Ii.GEE on photographic matters, and the painstaking typing of Mrs.Sheila KING. The writer is particularly indebted to Miss Margaret DOE for her considerable assistance in the final compilation of photographs and diagrams. Dr.D.V..LGER who supervised and stimulated the work is due special thanks. Research Studentship from the Natural Environment Research Council supported the study from 1963 to 1966 and is gratefully recognised. Finally, most particular thanks are due to my brother for his mechanical expertise with a Herculean field vehicle, and to my parents for their con - stnnt encouragement and support. 7 INTRODUCTION PREVIOUS RESEARCH It is now well over a century since PETZHOLDT (1843) published the first figures of a Rhaetian brachiopod, Spiriferina junfOrunnensis (now a nomen oblitum of Zugmayerella uncinata). Since then, although almost all the first descriptions have appeared in only two monographs, they have in - volved the first application of techniques which have revolutionised brachiopod systematics. ZUGMAYER, in 1880, described in detail how transverse parallel sections could be made using guide lines on the ventral and dorsal valves. His five transverse serial sections of Oxycolpella oxycolpos are the first known and together with those published by BITTER (1898) of another Rhaetian species, Carapezzia geyeri (see plate A herein), were not to be excelled in quality for several tens of years. ZUGM.iiYER illustrated the internal structure Bf each of the species he described with what he termed a 'Cardinal-Schlifft. This constantly oriented section lies close to the plane of articulation and was carefully chosen to reveal the maximum internal detail. The results he obtained enabled ZUGMAYEB. to make the first divisions of the catch-all genera Terebratula and Spiriferina. Similarly, calcination of the shell by burning, a technique commonly attributed to BUC}G'IAN (1918), who employed it to reveal muscle scars, was used to considerably better purpose by ZUGMLYER (1880) to accentuate the contrast of the internal elements of his ground specimens (see plate 8, fig.3 herein). However, it was SUESS, professor of geology in the University of Vienna and one of the leading scientists of the nineteenth century, who laid the foundation for the study of the alpine Rhaetian and its brachiopod fauna. Born in London, SUESS published his 'Brachiopoden der Kdssener Schichten' (1854) at the age of twenty three. His later papers with OPPEL (1856) on the equivalence of the intra and extra-alpine Rhaetian beds, and with MOJSISOVICS (1868) on the litho-faunal facies of the Rhaetian, are funda - mental. Remarkably, SUESSts wider interests led him to initiate the scheme 3 whereby Vienna today receives her water supply from the Alps, and to become a leading Liberal member of the Austrian Parliament (see GREGORY 1915). Prior to SUESS's monograph only three Rhaetian species had been des - cribed: Austrirhynchia cornigera and Calcirhynchia subrimosa by SCEthFHAUTL (1851), and Zugmayerella uncinata by the same author two years later. No further original descriptions followed for twenty six years until ZUGMAYER (1880) published the results of his work in the Piesting Tal region near Vienna. This area has a remarkably rich and diverse Rhaetian fauna and must inevitably have become a keystone in its study. ZUGMAY1!:;li, living in the Piesting Tal, was able to devote considerable time to collectir7.' This coupled with his advanced technique, discussed above, and the work of SUESS and MOJSISOVICS (1868) on the facies of the Rhaetian, enabled ZUGMAYER to produce a monograph including not only ten new species but in addition, a table of the facies distribution of the twenty six species then known. Ls a consequence of ZUGMiYER'S thorough study BITTNER felt able to leave the Rhaetian fauna out of his 'Brachiopoden der alpinen Trias' (1890). However, he included short descriptions and figures of specimens from the late Norian of Drnava (south govakia), which may be attributed to Rhaetian species. Similarly BITTNER's several papers on the Norian fauna of north east Turkey (1891, 1892a, 1895) involve Rhaetian species. Some seventy years

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