Bulletins of American paleontology (Bull. Am. paleontol.) Vol 301

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COM P ZOOU, LIBRARY MUS '^'& MAY 1978 HARVARD BULLETINS '^'^'-"''••^" OF AMERICAN PALEONTOLOGY (Founded 1895) Vol 73 No 301 THE BORING GTENOSTOMATE BRYOZOA: TAXONOMY AND PALEOBIOLOGY BASED ON CAVITIES IN CALCAREOUS SUBSTRATA By Robert A Pohowsky 1978 Paleontological Research Institution Ithaca, New York 14850 U S A PALEONTOLOGIGAL RESEARCH INSTITUTION 1976-1979 President Secretary Harold Duami _ _ - - _ - Vice-President Philip C „ Vokbs LeRot Wakelet Ernestine Q Wright Treasurer Director E O JLatherine V _ .„ Assistant Director Assistant Secretary, Assistant Treasurer Counsel - Representative AAAS - - W Palmer Peter R Hoovix _ - JIebecca S Hakkis Armand L Adams Richard G Osgood, Council Jr Trustees Katherine V W Palmer (Life) John Pojeta, Jr (1975-1978) Ruth G Browne (1976-1979) Kenneth E Caster (1975-1978) John Margaret Duane K Cisne (1976-1977) S Harris (Life) L Rebecca Heroy (1975-1978) LeRoy (1974-1977) B O William A Oliver, Norman Sachs, Jr (1974-1977) Daniel B Sass (1974-1977) Harold E Vokes (1975-1978) Philip C Wakelet (1976-1979) Ernestine Q Wright (1976-1979) Jr (1976-1979) BULLETINS OF AMERICAN PALEONTOLOGY and PALAEONTOGRAPHICA AMERICANA Katherine V W Palmer, Editor Peter Doris C Hoover R Brann Assistant Assistant Advisory Board Kenneth A Hans Kugler E Caster Jay Glenn Marks Myra Keen Complete titles and price list of separate available numbers may b« had on application For reprint, Vols 1-23, Bulletins of American Paleontology lee Kraus Reprint Corp., 16 East 46th St., New York, N.Y 10017 U.S.A For reprint, vol I, Palaeontographica Americana see Johnson Reprint CorNew York, N.Y 10003 U.S.A poration, 111 Fifth Ave., Subscription may price of $20.00 per be entered at any time by volume or year, with average Bulletins Numbers of Palaeontograpkica Ameri- volume for cana invoiced per issue Purchases ductible from income tax in U.S.A for professional purposes are de- For sale by Paleontological Research Institution 1259 Trumansburg Road New York 14850 Ithaca, U.S.A BULLETINS OF AMERICAN PALEONTOLOGY (Founded 1895) Vol 73 No 301 THE BORING CTENOSTOMATE BRYOZOA: TAXONOMY AND PALEOBIOLOGY BASED ON CAVITIES IN CALCL\REOUS SUBSTRATA By Robert A Pohowsky March 30, 1978 Paleontological Research Institution Ithaca, New York 14850 U S A Library of Congress Card Number: 78-53752 Printed in the United States of America Arnold Printing Corporation Ithaca New York 14850 CONTENTS Page Abstract — - Acknowledgments - _ 5 Introduction Prospects and objectives 11 Historical resume IS 20 Procedures and techniques — 22 Repositories Anatomy and biology of boring bryozoans 23 Interpretation and taxonomy of bryozoan borings 30 General considerations Glossary Morphology 31 - of bryozoan borings 35 42 Systematic paleontology 46 Ropalonariidae Nickles and Bassler, 1900 Orbignyoporidae, — fam n fam Voigtellidae, n 68 — Penetrantiidae Silen, 1946 Cookobryozoonidae, Spathiporidae, n Immergentiidae Silen, 77 95 _ Terebriporidae d'Orbigny, — 92 fam fam n 57 109 847 120 1946 129 Family uncertain Stratigraphic ranges of genera 135 Taxonomic 137 positions of species 140 Evolution and phylogeny Topics and further investigation .: 147 References 150 Plates 159 Index _ 183 LIST OF TEXT FIGURES Page Text-figure 1+ Ropalonaria venosa Ulrich Terebripora ramosa d'Orbigny A B C-E F Terebripora Spathiopora sp sp G Terebripora A Spat/iipora sertum Fischer sp 16 B-E F G H comma Spathipora (Soule) OrbignyoPora} capillarts (Dollfus) Orbignyopora archiaci (Fischer) Haimeina michelini (Terquem) Autozooids and stolons from genera A Penetrantia sp B Spathipora C D Orbignyopora archiaci (Fischer) Immergentia sp A Voigtella rcgalis, n gen., n sp B Marcusopora C Foraripora pesavis Voigt and Soule 26 sp 71 ripleyensis, n gen., n sp 136 Stratigraphic ranges of genera (13) Early astogeny boring Bryozoa in Al Immergentia A2 B3 Terebripora ramosa d'Orbigny Marcusopora ripleyensis, n gen., n sp Cookobryozoon lagaaiji, n gen., n sp Orbignyopora ? tridelta, n gen., n sp B4 Penetrantia B5 Spathipora elegans Fischer B6 Spathipora occidentalis, CI Ropalonaria venosa Ulrich C2 Haimeina michelini (Terquem) C3 Spathipora ambidextra, D Penetrantia Bl B2 sp sp ? sp n sp n sp _ l+Z THE BORING CTENOSTOMATE BRYOZOA: TAXONOMY AND PALEOBIOLOGY BASED ON CAVITIES IN CALCAREOUS SUBSTRATA Robert A Pohowsky ABSTRACT (Upper Ordovician—Recent) live immersed in calThe chemical means All careous substrata that the organism penetrates by purely boring Bryozoa known species are here assigned to the order Ctenostomata Busk, 1852, [? mknown »« anacluding Penctrantia Silen, 1946] Until now, the group has been parts of Holotomical detail almost exclusively from studies on the internal soft with various cene species Fossil remains of these bryozoans have been confused borings and shallow pits of diverse origin The classification of previously recognized species is revised, and 13 Casterogenera including 48 species are recognized There are six new genera: Voigtella pora Cookobryozoon, FischercUa, Marcusopora, Orbignyopora, and reservaCast'cropora and Fischerella are assigned to the Bryozoa with slight Cookobryozoon lagaaiji, Imtion The following 20 new species are erected: mergentia atypica, I boydekifia, lanceolata, losancjdina, I patagoniana, I subangulata, Marcusopora riplcyetisis, Orbignyopora? cornbrashica O.? tridelta S Penetrantia soulei, Ropalonaria? bugei, Spathipora ambidcxtra, S brevicauda, Voigcheethami, S magnhorticellopsis, S occidentalis, Terebripora miniaiura, and V secunda and figured material includes the ancestiulae and early astogenetic tella regalis, New reversed stages of several fossil species In addition, various heterozooids, several zooids and teratological abnormalities are recognized in fossils of genera Some groups are characterized by the occurrence of enantiomoiphic autozooids There is substantial evidence that at least the initial portions of entire colonies occur in dextral and sinistral forms in some species The confinement of zooids beneath an inflexible calcareous shield has sonnetimes been accompanied by the development of features resembling those which are generally associated with the cheilostomes The occurrence of ovicells in Spathipora cheethami, a boring species from the Middle Jurassic (Cornbrash) of England, is an example In Penctrantia densa Silen (Recent), a probable ctenostome, the presence of ovicells, an operculum, and a calcareous body wall suggest indirectly that the Cheilostomata may be polyphyletic Ovicells (and opercula?) may be more likely to develop in lineages wherein the zooids are protected by a solid external skeleton (of bryozoan or molluscan origin) The possibility of independent evolution of these features in at least two groups of "shielded" ctenostomes {i.e., encrusting in Penetrantia Silen and in one or more independent lines of calcified, ctenostomes) is worthy of thorough consideration ACKNOWLEDGMENTS by no means an individual effort; without the This study generous assistance of numerous people, its completion would have been impossible Unfortunately, space precludes individual mention is of am in some way indebted, and institutions may be cited as having everyone to persons whom I but the following been particularly helpful Many ment of sincere thanks are due, first to the faculty of the Departof the University of Cincinnati for instruction pre- Geology requisite for the undertaking of this study especially indebted to my advisor, Dr suggestions, enduring interest in the manuscript my In this regard, Kenneth E Caster, work, and critical I am for his reading of Bulletin 301 Much of the Museum British Keepers and to me research for this study was undertaken at the (Natural History), and I am grateful to the staff of that institution for the many my all, during visit I whose encouragement, my efforts The larly helpful: indebted, above assistance, to courtesies shown Miss P L Cook, and discussions greatly expedited members were also particu- following other staff P J Chimonides, R Cleevely, R Croucher, V K Din, R Harris, N R Wise am J Morris, P C Nuttall, R Symes, The valuable aid of Museum Microscopy Unit of the Electron Through the courtesy is J Taylor, and photographers and the staff also gratefully acknowledged of R V Melville, I obtained some im- portant specimens and photographs from the Institution of Geo- (London) His helpfulness, and that of A Horton and Mrs B Coleman (I.G.S.) is much appreciated logical Sciences The specimens examined during the course of this investigation are housed in the various repositories listed herein museum curators who diligently prepared To the many requested loans go my sincere thanks In a few cases, photographic negatives were sent in lieu of actual specimens, and I am grateful to E national d'Histoire naturelle, Paris) and Foundation, Los Angeles) for J Buge (Museum D Soule (Allan Hancock extremely helpful service The this thoughtful donation of specimens by the following persons greatly appreciated: W C Banta, B W Cameron, Miss P L A B Hastings, D P Najdin, F Strauch, and F Westphal and prepared slides also Cook, I am Marcus for sending me the bored studied by her late husband, Prof Dr especially indebted to Mrs shells is E Ernst Marcus Much of the literature on boring Bryozoa is not in English, and immensely helpful translations were prepared by J Cosby, Mrs F Vieira, Mrs H Sabo and Mrs J Sandborn Mrs Vieira's lengthy translation of Marcus, 1938, (Portuguese) was especially useful During the research phase of this work, some aspects of the investigation were discussed with Hillmer, D Jebram, R Cuffey, A B Hastings, G W J Kennedy, C T Scrutton, L Silen, J D and Mrs Soule, Prof E Voigt, and B Walter, in addition to some of the persons cited previously I thankfully acknowledge the beneficial information and advice offered by numerous people, but anv misconceptions or inaccuracies contained herein are strictly those of the author Boring Ctenostomate Bryozoa: Pohovvsky Research for this study was made possible and generously sup- ported by an N.D.E.A Graduate Fellowship at the University of Cincinnati and a Fulbright-Hays Grant from the U.S.-U.K Educational Commission in London INTRODUCTION The boring Bryozoa, here assigned to the order Ctenostomata Busk, 1852^, are an ethologically defined group of ecologically aberrant species which live totally "immersed" within solid calcareous substrata in the marine environment Except for the operculum in the genus, Penetrantia Silen, 1946, only the lophophore ever extends above the surface of the inhabited object (Of course, sexual reproduction involves the free-swimming larvae common to all bryoProbably because of their sheer abundance, the zoans.) gastropods and pelecypods are the substrata most ployed in shells of commonly em- post-Paleozoic seas, but other calcareous objects are also — especially in the Paleozoic, when brachiopods and crinmost readily available "hosts." (The term is placed in quotation marks because none of the bryozoans considered herein show any evidence of being parasitic; the boring habitus is solely a means of ensuring protection and anchorage while engaging in the invaded oids were the heterotrophic, filter-feeding process characteristic of bryozoans.) Boring bryozoans penetrate exclusively by chemical means and must be clearly differentiated from the relatively few species of Ctenostomata which reputedly employ mechanical processes to penetrate less substantial media These species and their typical substrata are listed below: Hypophorella expansa Ehlers, 1876; leathery tubes of polychaete annelids {e.g., Chaetopterus and Terebella) Watersian-a paessleri Calvet, 1912; tunic of colonial ascidian, Polyzoa gordiana Harmeriella terebrans Borg, 1940; calcareous walls of cheilo- stome Bryozoa {e.g., Tubiporella levinseni) Bulbella abscondita Braem, 1951; rotting wood Those Cheilostomata which are known to produce shallow pits on encrusted shells [e.g., Hippothoa divaricata Lamoroux, 1821; Electra monostachys (Busk), 1854, pi 10, fig 5] are also excluded from the group under consideration 'One of these, Penetrantia Silen may be a rheilostome Bulletin 301 Beyond question, the borings discussed the bulk of the known fossil work represent in this Ctenostomata Encrusting species pre- served ("immurated") beneath sessile benthonic organisms (e.g., Arachnidium brandesi Voigt, 1968) are eminently worthy of additional study but are known from comparatively few occurrences Putative fossil ctenostomes preserved as non-immurated encrustations {e.g.y Ascodictyon Nicholson and Etheridge, Eliasopora Bassler, and Marcusodictyon Bassler) lack well-documented Recent analogues within the order and are, therefore, of questionable affinity One such genus, Vinelloidea Canu, was recently recognized as a foraminifer (Voigt, 1973) As if by the pelecypods-bivalves-lamelli- refusing to be outdone branchs, and perhaps acquiring impetus from the recurrent flurries have been published concerning the of energetic perorations that proper name for the phylum Bryozoa, boring bryozoans have also been termed "burrowing," "penetrating," "perforating," and corresponding non-English equivalents thereof The group is united, fundamentally, by a common mode of life, and it may have arisen through the convergence of several lineages of epibenthonic ctenostomes But even monophyletic, its if assemblage were unquestionably this ethological informal nomenclature would clearly be without legal restraints, so an author powers of persometimes expedient to free to exercise his is sonal choice in the matter Moreover, it is use several different names alternatively within a given discussion in order to avert the monotony of repetition If the reader can under- stand what group of organisms fice If explicitly define the Within ous is being considered, any term will suf- nomenclatorial leeway exists, an author should briefly but group he is discussing this study, "boring" informal synonyms are bryozoans predominates, but variused also Brevity and alliteration generally vindicate the vapid and groundless pun For additional discussion Pohowsky (1974), which of standard zoological The present work consult concerned primarily with a revision of the Bryozoa; evolution of the group throughout — matters grounds for the preferred usage nomenclature for bryozoan borings is classification of all boring vician terminological of also gives Recent) Although the its it consequently deals with the entire known range (U Ordo- soft parts of these organisms are of Bulletin 301 182 Explanation of Plate 23 Page Figure 1-3 127 Immergentia spp Young colonies, showing budding pattern at ancestrula Stolons diverging from opposite sides of ancestrula growing (fortuitously?) to form obtuse angle, with ancestrula at apex On fragment of Ranella maiginata (Gmelin) Under D 52393 (on 83445) polish Pliocene Italv BMNH BMNH X Four colonies with stolons, without mature autozooid Arrows indicating ancestrulae which have just begun to produce stolons Largest colony (top, left) shows slight expansion of stolon perhaps representing developing autozooid Two colonies Larger one produced one autozooid (top, with paired lateral stolons arising slight^' distal center) to it Colony with ancestrula (arrow) and autozooids Paired stolons arising distal to autozooids near ancestrula 4-6 ; 27 125 Immergentia lanceoiata, n sp McHolotype On Polinices sp L Pliocene (Kalimnan) Donalds Bank, Muddy Creek, near Hamilton, Victoria, ; BMNH D 52383 (SEM G 39704-9, shell B) Surface of colony in shell cast in in comparison with / hoydckiiia men dry; X 23 Austr STUB 26), figs 5, (on BMNH Apertures large (cf PI 22, fig l) Speci- SEM of polyester cast of shell in fig Zooids similar to boydekina, distal end stouter, strongly flared in some specimens Bilaterally opposite principal stolons arising and numerous adventitious stolons zooids, distal to emanating from principal stolons; X 50 (approx.) / Another portion of cast shown in fig Autozooid at right producing adventitious stolon, this growing toward surface and fusing wih distal end of zooid in center of figure; 7-9 X 90 (approx.) Immergentia subangulata, n sp Crepidula sp Recent Bav of Santos, Brazil (Gift, Mrs D.52384 (SEM STUB 6, shell D) E Marcus.) Surface of colony in shell fragment cast in figs 8, Arrow corresponding to arrow in fig 8, indicating zooid in center of fig Specimen inked; X 25 On BMNH SEM of polyester cast of colony shown in fig Arrow similar to arrow in fig 7, indicating central zooid in fig Proximal end of zooids leaning proximally along stolon Zooids commonly flexed in that direction, laterally, or both Paired lateral stolons diverging distal to zooids near bottom of figure; X 42 Enlarged view of cast in fig 8, seen from opposite direction Central zooid (indicated by arrow in fig 8) producing adventitious stolon, this growing toward surface and fusing with distal end of zooid in foreground Small saccate bodies at upper right of alga! or fungal origin; 125 (approx.) X 125 Bull Amer Paleont., Vol 73 Plate 23 Bull Amer Paleont., Vol 73 Plate 24 Boring Ctenostomate Bryozoa: Pohowsky 183 Explanation of Plate 2+ Page Figure 127 Immergentia spp Surface of colony, resembling / boydekina and / subangulata Three of zooids sinistral Treated with ink; under polish In gastropod Colony labeled "Terebripora orbignyana Fisch." by Marcus Recent Bay of Santos, Brazil D 52394 (RAP 236) X 40 BMNH ; Immergentia atypica, 125 sp n Holotype Surface of best-preserved portion of colony showing bilaterally opposite branching of stolons, these just proximal to apparently monomorphic autozooids with nearly circular apertures situated medially along stolon Specimen dry In Maortcrypta sp Tertiary Lower Waipara Gorge, D.52385 (RAP 301) N.Z X 23 BMNH ; Immergentia losangelina, n 126 sp Holotype, showing surface of large and densely spaced zooids Apertures rarely separated by more than twice their length Specimen dry In Olivella biplicata Swainson U Pleistocene, Palos Verdes Sand Pacific and Oliver Streets, San D 52386 (on Pedro, Los Angeles Co., Calif G.72500-5) X 24 BMNH BMNH ; Immergentia californica Silen, 1946 In Tegula ligulata Menke Recent Tidal pool, Whites Point, 50 Calif (Negative court, of J D Soule.) Coll Soule; 122 X Immergentia patagonlana, n 126 sp Surface of holotype, showing profuse development of tubuadventitious stolons, strongly enantiomorphic zooids lets, About half of zooids dextral Zooids lying in shallow depressions indicated by differentially stained, deeper shell layers Treated with ink; under polish In BuccinanoPs D 52388 (on cochlidium Kiener Recent Argentina BMNH BMNH 1854.12.4.466) ; X 56 Immergentia spp 127 Syntype of Terebripora orbignyana Fischer, 1866, showing zooids (2 dextral, sinistral) in translucent shell Specimen dry In pelecypod Recent Arcachon, Gironde, France Di-0207-b (within ink circle on smaller of shells) MNHN X 31 ; INDEX NUMBER 301 Note: Light face type refers to page numbers Bold face type refers to plate number Australia Victoria abscondita, Bulbella AHF 128, 172, 173, 181, 182 Aetea 95 93, 124, 125, 18,50 (Allan Austria Hancock 22,112,177 108 75, 89, 166 Foundation) Alabama Dallas Co Alcippe 89, Voslau-Wien Bugula avicularia, 139 54 Alcyonidium B ambidextra, Spathipora 18 106, 107, 141, 143, 177 52, 161, 162 178 174, 181 Ampthill Clay Ancilla Ancillaria anglica, Graysonia angulata, Immergentia antillarum, 140 124, 139 Terebripora 137 antiqua, Entobia Terebripora arachne, Ropalonaria Ropalonaria? 78 128 127, 128, 181 147 Grund 137-139 67, 68, 137 Bascomella 139, 140 bassleri, Terebripora 138 Antwerp Docks berquisti, Graysonia biplicata, Olivella BMNH 162 138 8, 50 164 22, 29, 47, 52, 54-56, 59, 65, 67, 69, 86, 8991, 93, 96, 103- 52, 53, 109, 112, 114, 117 118, 124129 132, 160164, 167-169, archiaci, Orbignyopora 17, 26 27, 5863, 65-67, 138, 164 59, 60, 138 127, 183 108 Terebripora Argentina Santa Cruz argile de Dives ("vaches noires") argiles oxfordiennes de Dives arietina, Ditrupa "Arkona beds," lower shales of Hamilton Gp 56, 162 botellus, Ropalonaria Bowden Marl Bowerbankia 162 177 31, 55 Immergentia 22 124, 125, 181183 brandesi, Brazil Bay lie of Santos 89, 90, 105, 108, 109, 114, 117, 118, 125, 127, 129, 171, 175-177, 179 180, 182, 183 62,163 Ascodictyon attenuata, Rhopalonaria Ropalonaria 64, 139, 61 163 brevicauda, Spathipora 16 brevis, Penetrantia atypica, Immergentia 24 121 125, 183 171-183 49, 139 59, 164 boydekina, Arachnidium 52, 161, 169, 160 of Natural History) 138, 148, 161, archiaciana, Ostrea 47, (British Museum Arachnidium 129 140 183 Blanchester Mbr., Waynesville Fm 40 2,3 22,47.52-57, Terebripora 47, 57, 78, 88, (?) Belgium, 184 37,97,104 174, 175 79 84, 139 INDEX cochlear, 183 Buccinanops Pycnodonta bugei, Ropalonaria? Bugula 164 cochlidium, 47,55,56, 162 147 Buccinanops 183 comma, Spathipora 18 11 11 Bulbella Bundenbach Burgess Sh Burnupena 16, 17, 19, 27, 34, 37, 43, 51, 77, 81, 101, 111, 138, 144, 177 170 Terebripora 97, 101, 138, 177 concharum, Penetrantia 10 24, 39, 78, 79, 85, 87, 139, 94 California contorta, Terebripora contraria, Neptunia 93, 173 Kern Co Newport 101,177 122 Grove San Pedro, Los Angeles Co Pacific Conus Cookobryozoon 173 81, 120, 122, Ripley Cornbrash 139, 183 89, 105, 109, 118 112, 176, 178 cornbrashica, Gp Calyptraea Orbignyopora? 113, 174, 176 Camden Fra cancellata, capillacea, Fm 24 Caloosahatchee Fm Exogyra cucullata, Spathipora Cyclopuncta Cypraea Cyprus, Ayias, Cardiff Sh Mbr., 62, 132, 163, Cephalodiscus Chaetophorites Chaetopterus Chalk cheethami, Spathipora 150 55 46, 52, 130 danica, Iramena delalandi, Burnupena Delaware, Bethany Beach, Rehoboth delta, Liostrea 15 37, 81, 96, 99 65, 164 78, 87, 140 170 100, 175 162 dendriformis, 174 81,99,174 112, 178 169, 175, 181 32, 33, 129, 131-133, 137, 138, 163, 173 103, 104, 145 Chile Arica Cincinnatian Series 99 100, 138 58, 61, 131 Simeous 173 14 164 55 23, 93 166 59, Cryptopolyzoon Cucullaea 138 Casteropora 139 Candia 17 18, 61, Marcellus Sh 166 176, 182 Cryptomonas 17, 64 37, 58, 61, 6365, 138, 139 163 Terebripora 66-68, 164 166, 168 cretacea, Entobia Crete, Bythmia, 61 Crepidula capillaris, Orbignyopora Orbignyopora? 75, 89, 118 119, 166, 168 37, 67, 103, 164, 174 costata, Exogyra Crassetellites 178 62, 163 166 Terebripora 178, 181 14 40, 41, 92-94, Coon Creek Mbr., californica, Immergentia 169 138 169 106, 135, 141, 143, 148, 172, 105,126,176, 183 183 Whites Point 13, 19, 48 Ropalonaria Denmark, Fakse dcnsa, Penetrantia 11,12 18$ 139 78 9,28,29,31, INDEX Suffolk 32, 36, 38, 39, 78, 79, 81-85, 87, 88, 92, 139, 144, 146 168, 170, 171 Terebripora dilatata, Gryphaea discus, Microcyclus distans, Hippothoa Ditrupa ditrupae, Terebripora 52 Thorpe, St Andrew, Norwich, Norfolk Thrapston, Northamptonshire Yorkshire Entobia eocenica, Terebripora 138 161,162 163 18 169,177 Europe Exogyra 103, 174 90 137-139 138 92 166, 168 19, expansa, 87-89,96 138 169 177 Hypophorella 7, 27, 37 ditrupiae, 87 177 54 Terebripora divaricata, Hippothoa draconis, Polinices duplex, Alcyonidium falunica, Tere- bripora .19, 20, 21 15, 19, 41, 66, 111, 114-118, 138, 178-180 E Farella Ficus Fischerella echinicola, 138 176 7,29,148, 169 59, 61, Terebripora Echinophoria Electra fischeri, Terebripora elegans, Spathipora 15 27, 37, 81, 96 elongata, Terebripora Florida 178 176 floridana, Bedfordshire Bournemouth Flustrellidra Foraripora "Formengruppe C" 67, 103 Chicksgrove, Tisbury Chideock, Bridport Easton, Isle of Dorset, Dorset Faringdon Helpstone Long Crendon, Buckinghamshire Lyme Regis, Dorset Mickleton, Gloucestershire 108 91 flasser Zone 61, 62, 131, 163, 173 France, Antibes, Alpes 108 88 67,103 Maritimes 109, 128 Arcachon, Gironde 121, 128, 183 Barneville-sur- 108 90 Mer, Manche Brassempory, 62 Landes "Curcey" 90, 167 59,164 91 108 Dax 67, 103, 164 174 52 Dives-sur-Mer, Calvados 52,56,161 162 Stockton, Warwickshire Stroud Gloucestershire 60, 70, 129, 130, 137, 140 131, 133 Fossil-Bank "a", Seifen H, Wild- Peterborough, 176 80 Calyptraea 19,138,149 78,92,104 52,103,161 162 Terebripora England probably 109, 112, 118, Harney Pond Canal, 89,105,118 Glades Co eltaninae, Northamptonshire 116, 117, 138, 179 99, 107, 112, 138, 141, 143 145 174 178 59, 60, 138, 164 Eliasopora 31 107, 176, 177 129, 134, 135 137, 139, 166 Gan, BassesPyrenees Hettange, Moselle 90 90, 167, 168 186 59 91 INDEX GPIUB Manthelan, Indre109, 114, 116, 128, 175, 178, 180 128 114, 178 108, 128 67,68 97 Lagus Saubrigues, Landes LePoivre, Bordeaux Luc Marseille (Geol.-Pal Inst Univ Bonn) 92, 104, 108 et-Loire gracilis, Bowerbankia graphicus, Ropalonaria Graysonia Grunder Schichten? GSM (Inst Geol Sci., London) Touraine 22,52,161, 162 90 Orientales Pont-le-Voy, Loire-et-Cher Saint-Germain-leFouilloux, Mayenne 97, 109, H 118 132 Haimeina 8, 17, 38, 39, 81, 88, 90-92, 135, 138, 141, 143, 149, 167, 89, 105, 109, 118, 127, 128, 169, 175, 181 Trouville-sur-Mer, Calvados Hamilton Gp Calvados 62, 63, 163, 173 52, 161 fusiformis, Bascomella 139 Harmeriella Hawaii, Haena Bay, Kauai Keaukaha, Hilo helicoides, Priamus G 90 163 91 Terebripora Hippothoa Hudson River Gp Hungary, Delinyest, Krasso 173 hyeroglyphica, Terebripora 130 Marmorgraben bei Mittenwald Seifen bei Dierdorf 62, 131, Stuttgart-Degerloch Unkel-MUhle bei Eitorf/Sieg Georgia, Lumkin, 138 7,18,49 48 128 138 37 Hypophorella Stewart Co 7, 27, 89 139 gigantea, Bascomella giganteum, Plagiostoma I 167, 168 Immergentia (Geol Inst Univ Koln) 22, 23, 24 22,173 givetiana, Ropalonaria glandiformis monstrosa, 139 Ancillaria (Geol Landes Nord Westf.) 181 22, 61, 62, 131, 163, 173 40 163, 173 gosaviensis, 9, 15, 17, 19, 23, 25, 26, 30, 32, 40, 42, 57, 81, 110, 112, 116, 117, 120127, 132, 137141 143, 149, GLNW gordiana, Polyzoa 181 140 hieroglyphica, Lagerdorf, Holstein Glosselettia 86 124 Heteronema Germany, Globigerina 106, 139, 145, 168, 171 52 Villers-sur-Mer, Penetrantia 139 140 89 161,162,165 Gryphaea Perpignan, Pyrenees GIK 74 165 31, 55 22, 73, 181-183 Immergentia? incerta, Spathipora 42 108, 138 Indiana, Hanover Versailles 48 48 indistincta, Spathipora 78, 87, 88, 139 inflata, Ancillaria 187 19, 138 174, 181 INDEX laxa, Spatipora intermedia, 176 134, 166 78,87, 140 Echinophoria Iowa, Keokuk Iramena 85, 87, 139, Ropalonaria Terebripora 144 139 137 59,88, 121, 127, 128, 164, Italy 182 109 128 129 Artigiana "Astigiana" Asti, Tubiporella Piedmont 183 162 Tegula Liostrea Penetrantia lissajousi, Immergentia? Lithuania, Popilani longicauda, Spathipora 177 longirima, Spathipora 16 37,99,100, 103, 138, 175 128 89 128 118 Castel Arquato 42, 139 52, 161, 162 100, 108, 138, 89, 109, 121, Bordighera 138 169 levinseni, ligulata, irregularis, 87, 88, Leporemax losangelina, Immergentia Lower Portland lutheri, 89, 105, 107, 24 126 183 beds 108 55 Monochrysis 176 177 106, 129 176 Colli Astesi Colline Sensi Orciana, Toscana Piacenza Piedmont 109 129 127 181 89,109 129 129 89, 105, 128, 176 Jamaica, Bowden 59,164 17 106, 107, 176 59-61, 138 183 62, 132, 163 173 Marcusodictyon Marcusopora K keokukensis, 134, 135, 139, Fischerella Rhopalonaria Ropalonaria Keokuk Ls Kimmeridge Clay Kossener Schichten 166 134 139 134, 166 52,162 90 marginata, Ranella Marlstone marshii, Ostrea Cookobryozoon 13,14 lagaaiji 41,92-94, 141, 143 148 172 173 Rhopalonaria Ropalonaria Mediterranean Terebripora 61 64, 97, San Antonio, 84 California 85 MGU (Moscow State 22,69,165 University) michelini, Haimeina 8, 17, 38, 39, 81, 90-92, 106 138, 139, 141, 143, 145, 149, 167 168 171 23 125, 181, 182 138 139 101 Ibiza lanceolata, Immergentia 161 11 medialis, 61 139, 163 latesulcata, 57, 66, 70, 118-120, 137 141, 143, 166 164, 182 90, 167, 168 Mazon Creek lambtonensis, Rhopalonaria Ropalonaria Baleares Mexico, San Benito Islands, Baja L 140 magnivorticellopsis, Spathipora manzonii, Terebripora Maoricrypta Marcellus Sh Val d'Andona, Piedmont M magna, Heteronema Vioa? Microcyclus 188 90.137,139 163 INDEX Ohio, miniatura, 20 Terebripora Acton Lake, Hueston 15, 111, 115- Woods 117, 175, 179, MNHN Monobryozoon Monochrysis monostachys, Electra mucronata, B., 47, 48, 160 47, 51 Cincinnati Clarksville (Mus nat Hist 22,52,59,67, nat, Paris) 99, 112-115, 121, 127, 161, 164, 174, 178, State Park, near Oxford 180 47,160 48 48 Oregonia Waynesville Olcese Sand Mbr., Fm Temblor 183 93 Olivella 55 Ontario 93, 173 183 63 62, 163 Thedford operculata, 69 176 Zone of mutabilis, Nassa 36, 86, 139 Penetrantia orbignyana, Terebripora 114, 116, 121, 127, 128, 138, 178, 183 17, 26, 27, 45, 57-62, 64-67, 135, 138, 164 N Orbignyopora 176 169 Nassa Neptunia S New 169 58,63,64, 133 62 Walcheren York 138, 139, 141, 143, 163, 164 86, Hubbardsville Lockport 61,62,163 62,132,163, 173 87 Morrisville New Zealand P pachyderma, Terebripora Hawkes Bay Lower Waipara 105, 117, 118, 138, 176 90 paessleri, Gorge Otago Peninsula, Little Papanui Moko Hinau Island, Hauraki Gulf Slipper Island 125,183 85 Watersiana Palos Verdes Sand Parasmittina parva, Penetrantia Immergentia Umuahia Pecten Penetrantia 126, 183 84, 80 139 .16,17 105-107,141, 143, 175, 176 24 121, 122, 126, 183 24 7,17,23-33, 36, 38, 39, 43, 77-89, 92, 99, 102-104, 135, 138-141, 143147, 149, 168171, 176 permiana, Rhopalonaria Ropalonaria occidentalis, 117,138 9,10,11,12 118 88 68,77 Terebripora patagoniana, Wanganui, North North America Nygmites parvicella, 84 123 89,90,129 Island NHR (Naturh Riksmus., 22,82,84, Stockholm) 169, 170, 171 Nigeria, Eastern, Uncinulus 40 Globigerina Maraekakaho, oehlerti, 161,164 Ostrea pacifica, "Fossil Creek", Spathipora 4,5 37,60,62-68, Orbignyopora? Netherlands, Kaloot, 77 74,75,77, 139 165 pertenuis, Ropalonaria 133 189 19,49,139 INDEX pesavis, Foraripora 60, 70, 129, 130, 140 19, 36, 49, 50 54, 108, 110^ 123 117, 124, 137, 139, 141, 143, 177, 178, 180 Philippine Islands, Zamboanga Zamboanga City, philippinensis, Immergentia Ranella RAP (Robert A Pohowsky reference 112, 117, 122-124, 139, 149 9, 15, 19, numbers) 29, 148 92, 162, 164, 167, 168, 174 172, 173, 177, pilosa, Electra Plagiostoma Polinices 22, 59, 62, 69, 75, 82, 89-91, 100, 108, 109, 112, 114, 117119, 124, 125, 127-129, 132, 160, 163-166, 168, 170, 171, 173-175, 177181, 183 182 178 Polyzoa Porphyria portlandensis, Pyriporopsis 37, 92, 104, Portland Stone 145 108 rc^alis 72, 73, 75-77, 138 131 Priamus prima, Spathipora 138, 165 42 Terebripora Rhopalonaria 138 42, Stomatopora 138 135 49, 139 163 91 Nygmites Rhopalonaria? 134, 163 46, 53 Richmond Gp 47,48,160 119, 166, 168 Marcusopora 72,77,139 Pyriporopsis 37, 92, 104, 145 61,62,163 Museum) Ropalonaria quenstedti, Terebripora 90, 22, 62, R 174 radiciformis, Terebripora Rafinesquina ramosa, Terebripora 138 160 19 163 14,15,17-20, 37, 40, 45-54, 56, 58, 63, 64, 66 68 69, 74, 75, 77, 111, 134, 135, 138, 141, 143-145, 150, 160, 163, 138 radians, Calyptraea 70, 118-120, 141, 143, 166 61 62 64, 139, 163 19, 138 Rhopalonaria Ropalonaria Terebripora Rochester Ls ROM (Royal Ontario 137 164 Pycnodonta robusta, pusilla, Terebripora 162 75, 89, 118, ripleyensis, 77 Talpina 137 46, 47, 58, 61, 62, 68, 73, 77, rigidum, Plagiostoma Ripley Fm provularia, Schizophoria Psilonotenkalk pungens, 165 31 140 107, 176, 177 repens, Farella Vinella reticulata, Ficus reticulum, 69 72, 77, Proboscidactyla producta, Terebripora propinqua, Terebripora Protulophila proutana, 38,39,68-70, Voigtella portlocki, Terebripora? 164, 182 165 Ropalonaria? 2,3 22,52-57, 138, 139, 148, 161, 162 9, 14, 15, 17- 190 INDEX Rumania, "Lapugy", 89,108,118, Transylvania 124, 128, 174, 181 S Mary's Fm Santa Cruz Fm Schizophoria "2nd Mediterranean Stufe" secunda, 177 108 163 St 128 69, 71, 74-77, 166 Voigtella septemradiatus, 24 59 Pecten Serpula sertum, Spathipora ^ 18 16,17,96-99, 107, 108, 135, 138, 175, 177 89,96,98, 169, 177 Shetland spirulea, Serpula, Zone of Terebripora Stomatopora Tegula I38'i39 183 .^ Temblor Fm 9'5, sileni, 89, 109, 85, Penetrantia 89 128 139 sinefilum, 117, 138 Terebripora Skaneateles Sh Solnhofen Soule Collection 62 11 183 soulei, 10 Penetrantia Camden Dave Weeks 62, Ibd Place, Coon Creek, McNairy Co 75, 89, 119 166, 168 tenuis, Waccamaw Horry Co J 9' 1^' l^- 85-87, 144, ! 82 River, 100, 175 pora 15,16,17,18 10,15-17,19, 183 Terebripora (?) Timor Noil 104, 106-108, 111,112,131, J^' 40, 41 45-46! 49, 50, 52-54, 57-61, 66-68, 77 87-90, 9597, 101, 103, 105, 108-121, 124, 127, 128, 131, 137-139, 141, 143-145, 148, 149, 164, 169, 175-180, 82, 170, 171 26, 27, 32-34, 37, 43, 45, 51, 58, 61, 68, 69, 72, 77, 78, 81, 88, 89, 95- •>«> 163 139 138 ' Harmeriella Terebripora 19,20,21 Spathi- Boewan 47,57,88 74 73, 129, 165 timorensis, Rhopalonaria Ropalonaria 135-138, 141, 143-145, 165, Voigtella? 174-178 Spatipora 61, 64, terebrans, 169 Cape of Good Hope OA Chaetophorites Rhopalonaria Ropalonaria Terebripora Terebella South Africa, Port Nolloth South Carolina, 17^ 118 Tennessee 78,101,138 Palermo 138 139 160 T T^^P^"^ Sicily, Buccheri 49, 23 125,182,183 Immergentia 175, 180 subrotunda, Venus subsphaerica, 139 Bascomella 123, 139 suecica, Immergentia 123, 139 suecia, Immergentia Sweden, Flatholmen, 85, 123, 169 GuUmars Fjord 59, 109 Syria, Latakia 78,88,89, 101, 103, 138 Terebripora? 164 Streptelasma subangulata, sibirica, Spathipora 59, stellifora, 77,87,88,95 191 73 74, 139 69,72-74,76, 165 INDEX venosa, tridelta, Orbignyopora? Ropalonaria .5 58, 60, 65, 66, 141, 143, 164 14, 15, 17, 18, 37, 46-54, 56, 163, 173 138, 141, 143, 150, 160 triquetra, Glosselettia 7' Tubiporella Turkey venosum, Rhopalonaria Venus 114 "Saaret" (Sayaret), Lycia 112,178 128 Zengen 47 175,180 vesicularis, Gryphaea 165 vetusta, U Casteropora UCGM (Univ Cincinnati Geol Mus.) 22, 47, 52, 62, Terebripora Vinella Vinelloidea 69, 75, 89, 91, 100, 119, 132, 160, 163, 165, 166, 168, 173, Vioa? Beulahland, King William Co Bowlers Wharf, Middlesex Co 175 139 140 133 78 Vinella Uncinulus United States "upper Graywacke of Manche, graywacke with Spirifer" USGS (United States 62 Geological Survey) (National 22, USNM Museum U.S.S.R Chvalyusk, Saratov Province 22, 48, 57, 59, 61, 62, 64, 100, 105, 134, 135, 160, 163, 164, 166, 175- 177 78,88 69 52 "Kosminsk" Mt Chodsa Kasian, 100 177 109,128 Yorktown Voigtella 6,7 38.39.68-70, 72-77, 135 138, 139, 165 166 Voigtella? 69, 72-74, 76, 139 165 105 of Natural History) 90, 140 137, 139 Virginia, ulrichi, Ropalonaria 14 131-133, 138, 163, 173 131, 138 Voigt Photo Catalogue of Vorticella vulgaris, Cucullaea 130 106 166 W Waccamaw Marl Watersiana Waynesville 100, 175 Fm 47, 48, "Wurmlocher" 160 90 Y S W Schaartus, Tadschikistan 89, 101 Yorktown Fm 100, 109, 128 Sengiley, Ulia- novsk Province Trans-Caspian Province 69, 165 69, 165 Vol'sk, (Saratov Province) uva, Valkeria zelandica, Immergentia 165 27, Immergentia 123, 139 ZGM V (Central Geol Mus., Leningrad) vadosus, 22, 89, 101, 102 Crassetellites Valkeria varians, Terebripora 123, 139 minuta, 37 19, 166 27,37 138, 149 Zone of Serpula spirulea Zone of B mucronata 192 59,164 69 (No8 233, LII New 387 pp., 43 pis 236) „ 18.00 Zealand forams, Stroraatoporoidea, Indo-Pacific, Mio- cene-Pliocene California forams LIII 488 pp., 45 pis (Nos 237-238) Venezuela Bryozoa, Kinderhookian Brachiopods LIV (Nos 239-245) 510 pp., 50 pis „ _ 20.00 Dominican ostracodes, Lepidocyclina, moUusks LV LVI LVII LIX ™ (Nos 246-247) 657 pp., 60 r is Cenozoic corals, Trinidad Neogene mollusks 572 pp., 49 pis (Nos 248-254) Forams, North Carolina fossils, coral types, Cenozoic Echinoids, Cretaceous Radiolaria, Cymatiid gastropods 20.00 321 (Nos 255-256) Jurassic ammonites 20.00 ^ LVIII pp., 62 pis 18.00 ™ 18.00 „ 20.00 335 pp., 68 pis (Nos 264-267) Radiolaria, cirripeds, Bryozoa, palynology LXI „.„ (Nos 268-270) 365 pp., 31 pis Mollusks, Murex catalogue, Cretaceous Radiolaria LXII 375 pp., 44 pis (Nos 271-274) Trace fossils, ammonoids, 20.00 20.00 „ Silicoflagellates, raicrofauna 20.00 „ 320 pp., 56 pis (Nos 275-277) Chitinozoa, Spumellariina, Mexican Ammonites LXIII 20.00 305 pp., 39 pis (Nos 257-262) Cretaceous Radiolaria and Forams, Pacific Silicoflagellates, North American Cystoidea, Cyclonema, Vasum 314 pp (No 263) Bibliography of Cenozoic Echinoidea LX 20.00 — LXIV „ -_ 20.00 337 pp., 41 pis (Nos 278-281) Palynology, corals, echinoderms, Foraminifera, and crinoids LXV 20.00 687 pp., 49 pis (No 282) Ostracode Symposium (Nos 283-286) 639 pp., 62 pis LXVI 20.00 Crinoids, gastropods, corals, ostracodes LXVII LXVIII LXIX LXX 20.00 (No 291) Bryozoa 20.00 222 (No 295) 379 Paleocene Nigeria _ _ LXXII 20.00 233 pp., 28 pis (Nos 288-290) Paracrinoidea, ostracodes, cirripeds pp., 51 pis (Nos 292-294) 494 pp., 42 pis Turrids, gastropods, forams LXXI — „ 456 pp., 60 pis (No 287) Misc Paleozoic pp., 22.00 „ 20.00 _ 50 pis (Nos 296-300) 552 pp., 34 pis Crinoids, barnacles, forams, Stanford catalogue 20.00 Palaeontographica Americana Volume II See Johnson Reprint Corporation, 111 Fifth Ave., New York, N Y 10003 Monographs of Areas, Lutetia, rudistids and venerids (Nos 6-12) 531 pp., 37 pis Heliophyllum halli, Tertiary turrids Neocene Paleozoic cephalopods Fasciolarias, sponges 25.00 Spondyli, III (Nos 13-25) 513 pp., 61 pis Cephalopods, siphonophores, Devonian fish, gastropods, crinoids, Cretaceous jellyfish, brachiopods, bivalves 30.00 IV (Nos 26-33) 30.00 492 Devonian lycopods, brachiopods V VI Ordovician eurypterids, moUuskt, (Nos 34-47) 445 pp., 101 pis Pelecypods, Cretaceous Gulf Coastal forams 32.00 (Nos 38-41) _ 444 pp., 83 pis Pennsylvanian plants, Venericardia, Carboniferous crinoids, Trace Vn pp., 72 pis 40.00 fossils (Nos 42-46) 499 pp., 79 pis 45.00 Trilobites, rudists, crinoids, gastropods Vni (Nos 47-50) 337 pp., 36 pis Gastropoda, Devonian plants, brachiopods, Bivalvia , J7.50 BULLETINS OF AMERICAN PALEONTOLOGY VoL See Kraus Reprint Corp., 16 East 46th St., N Y 10017 U.S.A 334 pp., 27 pis (Nos 80-87) Mainly Paleozoic faunas and Tertiary Mollusca I-XXIII XXIV XXV XXVI XXVIII XXIX 306 pp., 30 pis (Nos 88-94B) Paleozoic, Mesozoic, and Miocene fossils (Nos 95-100) Florida Recent, 420 pp., Texas 58 XXXI XXXII XXXIII 12.00 _ 14.00 „ pis and South America 16.00 Cretaceous, fossils (Nos 101-108) 376 pp., 36 pis Tertiary mollusks, Paleozoic Venezuela, Devonian 16.00 fish (Nos 109-114) 412 pp., 34 pis Paleozoic cephalopods Cretaceous Eocene, forams 18.00 (Nos 115-116) 18.00 738 pp., 52 pis Bowden forams and Ordovician XXX York, _ Cenozoic XXVII New cephalopods _ (No 117) 563 pp., 65 pis Jackson Eocene Mollusks (Nos 118-128) 458 pp., 27 pis Mollusks, crinoids, corals, forams, Cuban localities (Nos 129-133) 294 pp., 39 pis „ Silurian cephalopods, crinoids, Tertiary forams, Mytilarca (Nos 134-139) 448 pp., 51 pis Devonian annelids Tertiary mollusks, Ecuadoran strati- 20.00 18.00 18.00 18.00 graphy paleontology XXXIV XXXV XXXVI XXXVII XXXVIII XXXIX XL XLI (Nos 140-145) 400 pp., 19 pis Forams, cephalopods, ostracods, conularid bibliography 18.00 (Nos 146-154) 386 pp., 31 pis Forams, cephalopods, mollusks, ostracods 18.00 (Nos 155-160) Forams, Eocene (Nos 161-164) 412 pp., 53 fish, rudists pis 18.00 486 pp., 37 pis _ Cretaceous rudists, Foraminifera, Stromatoporoidea (Nos 165-176) 447 pp., 53 pis Forams, ostracods, mollusks, Carriacot, „ 18.00 18.0« fossil plants (Nos 177-183) 448 pp., 36 pis South American forams, Panama Caribbean mollusks 16.00 996 pp., pi (No 184) Type and Figured Specimens P.R.I 18.0« _ „ (Nos 185-192) 381 pp., 35 pis „ Forams, mollusks, carpoids, Corry Sandstone 673 pp., 48 pis _ (No 193) Venezuelan Cenozoic gastropods 16.00 XLIII (Nos 194-198) 427 pp., 29 pis _„ Ordovician stromatoporoids, Indo-Pacific camerinids, Misissippian forams, Cuban rudists 16.00 XLIV (Nos 199-203) 365 pp., 68 pis Puerto Rican, Antarctic, New Zealand forams, Lepidocyclina, Eumalacostraca 16.06 (No 204) 564 pp., 63 pis Venezuela Cenozoic pelecypods 18.00 (Nos 205-211) 419 pp., 70 pis Forams, Crustacea, brachipods Recent mollusks 584 pp., 83 pis (Nos 212-217) „ Forams, mollusks, polychaetes, ammonites 16.00 XLVIII 1058 pp., pis (No 218) „ Catalogue of the Paleocene and Eocene Mollusca of the Southern and Eastern United States 18.00 XLIX (Nos 219-224) 671 pp., 83 pis Pencroplid and Australian forams North American carpoids South Dakota palynology, Venezuelan Miocene mol- 18.00 XLII XLV XLVI XLVII 18.00 18.00 luska, Foluta L (No 225-230) 518 pp., 42 pis Florida cirripeds, forams, Camerina, Ordovician conodonts Venezuela, LI 420 pp., 10 pis Antarctic bivalves, Bivalvia catalogue (Not 231-232) 1I.0» Linnaean Olives, 18.00 ... Marks Myra Keen Complete titles and price list of separate available numbers may b« had on application For reprint, Vols 1-23, Bulletins of American Paleontology lee Kraus Reprint Corp., 16 East... Research Institution 1259 Trumansburg Road New York 14850 Ithaca, U.S.A BULLETINS OF AMERICAN PALEONTOLOGY (Founded 1895) Vol 73 No 301 THE BORING CTENOSTOMATE BRYOZOA: TAXONOMY AND PALEOBIOLOGY BASED... bryozoans have an impor- tance far out of proportion to their ecological and economic Because of their peculiar mode of fossil life, records of the soft parts of any invertebrate organisms over,
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