&r^ BULLETINS OF AMERICAN PALEONTOLOGY (Founded 1895) MUS COMP 200U UNiVERS/TY ^^~^"~ No 291 A Lewis G Weeks Publication GENERIC REVISION AND SKELETAL MORPHOLOGY OF SOME CERIOPORID CYCLOSTOMES (BRYOZOA) By Osborne Barr Nye, Jr 1976 Paleontological Research Institution Ithaca, New York 148S0, U.S.A PALEONTOLOGIGAL RESEARCH INSTITUTION 1975-1976 President Harold Vice-President Duane Philip C Secretary Vokes LeRoy Wakeley Katherine V W Palmer Director, Treasurer David Assistant Director W Kirtley Rebecca Assistant Secretary, Assistant Treasurer Armand Counsel Representative E O AAAS S Harris L Adams Richard G Osgood, Council Jr Trustees Katherine V W Palmer (Life) John Pojeta, Jr (1975-1978) Casper Rappenecker (1973-1976) Ruth G Browne (1974-1976) Kenneth E Caster (1975-1978) Merrill Rebecca W Haas Margaret (1973-1976) Duane (1974-1977) LeRoy (1974-1977) O Axel A Olsson Norman Sachs, Jr (1974-1977) Daniel B Sass (1974-1977) Harold E Yokes (1975-1978) Heroy (1975-1978) B W Kirtley David K Harris (Life) S Philip C Wakeley (1973-1976) (Life) BULLETINS OF AMERICAN PALEONTOLOGY and PALAEONTOGRAPHICA AMERICANA Katherine V W Palmer, Editor Doris C Brann, Assistant Advisory Board Kenneth A Hans Kugler E Caster Jay Glenn Marks Myra Keen Axel A Olsson Complete titles and price list of separate available numbers may be had on application For reprint, Vols 1-23, Bulletins of Kraus Reprint Corp., For reprint, vol I, Subscription may St., American Paleontology New see York, N.Y 10017 U.S.A Palaeontographica Americana see Johnson Reprint Cor- poration, 111 Fifth Ave., price of $20.00 per 16 East 46th New York, N.Y 10003 U.S.A be entered at any time by volume or year, with average volume for Bulletins cana invoiced per issue Purchases ductible from income tax Numbers of Palaeontographica in U.S.A for professional For sale by Paleontological Research Institution 1259 Trumansburg Road New York 14850 Ithaca, U.S.A Ameri- purposes are de- BULLETINS OF AMERICAN PALEONTOLOGY (Founded 1895) 69 Vol No 291 A Lewis G Weeks Publication GENERIC REVISION AND SKELETAL MORPHOLOGY OF SOME CERIOPORID CYCLOSTOMES (BRYOZOA) By Osborne Barr Nye, February 12, Jr 1976 Paleontological Research Institution Ithaca, New York 14850, U.S.A Library of Congress Card Number: 75-^^572 Printed in the United States of America Arnold Printing Corporation Ithaca, N.Y CONTENTS Page Abstract Acknowledgments Abbreviations for repositories Introduction Taxonomic basis and procedure Approach Genera included 10 Synonymies 10 Generic diagnoses Techniques Biometrics Skeletal IS morphology 18 Zooecial wall structure 18 Microstructure - Orally oblique lamination Orally acute lamination Variation in thickness 18 21 23 23 Diaphragms and simple external walls 24 24 Introduction Interzooidal pores Zoarial brood chambers 38 Systematic descriptions 49 References cited 162 Plates 169 Index 222 ILLUSTRATIONS Text-figures 1-20 TABLES Tables 1-30 GENERIC REVISION AND SKELETAL MORPHOLOGY OF SOME CERIOPORID CYCLOSTOMES (BRYOZOA) Osborne Barr Nye, Jr Syracuse University ABSTRACT Thirteen post-Paleozoic cerioporid (Bryozoa) genera including 14 species have been restudied utilizing internal characters This approach applied to routine studies of Paleozoic tubular Bryozoa has produced relatively consistent taxonomic schemes Earlier studies of cyclostomatous Bryozoa were based on a relatively few, primarily external characters Variations of these characters generally reflect non-genetic factors The discovery of many new internal characters in post-Paleozoic cyclostomes expands the basis from which new taxonomies can be constructed and evolutionary inferences made Presumably as biological relationships of internal and external structures become known, estimates of genetic and non-genetic factors which control their variation will improve Genera were diagnosed on the basis of characters associated with zoarial growth patterns, microstructure of the zooecial wall, and occurrence of diaphragms Brood chambers, which are primary zoarial structures in the cerioporids studied, are too poorly known at present to provide taxonomic characters in supra-specific categories Cerioporids studied have ramose, massive, or frondose zoaria Ramose habit was produced by: (1) the formation of an axial endozone composed of nearly parallel growing, thin-walled zooecia which eventually bend radially and become thick-walled in the exozone (2) essentially like (1) as modified by a spiral budding pattern; (3) like (1), but zooecia stop growing orally after emplacement of frontal walls bearing peristomes; (4) repetitive hemispheric extensions of the basal layer to form an axial support structure upon which zooecia are initially adnate; (5) repetitive overgrowth in which each growth phase is composed of radially directed zooecia; (6) parallel growth of autozooecia which open only at growing tips Frondose habit is produced by bifoliate budding from a median layer Massive habit is produced by radial growth of zooecia Overgrowth and intrazoaria] anastomosis of growing branches are important modifications of growth habit in some genera Basal, intermediate, and terminal diaphragms; and simple external walls with restricted apertures can be identified in cerioporids They can be distinguished on their position within the zooecium, direction in which laminae flex when merging with the zooecial wall, occurrence of pseudopores, and occurrence of peristomes Basal, and perhaps intermediate, diaphragms formed floors to living chambers; terminal diaphragms presumably functioned as protective cover-plates to zooids in degenerative phases; simple external walls may have functioned as protective cover-plates by restricting the skeletal aperture to a small opening (peristome), through which feeding organs (the lophophore) had access to sea water Basal diaphragms were secreted by membranes on the oral side of the diaphragm Intermediate, terminal, and simple external walls were secreted by membranes on their aboral sides The secretion of intermediate, terminal, and simple external walls is related to the connection of interzooidal tissue through interzooidal pores Increased circulation through interzooidal pores, not possessed by most Paleozoic Bryozoa, may provide an adaptive advantage to most post-Paleozoic Bryozoa Observations of zooecial wall structure in cerioporids supports the "double wall" mode of growth model proposed by Borg (1926b, 1933) and expanded by Boardman and Cheetham (1969) In cerioporids, two major kinds of laminar structure can be distinguished In one group, laminae arch orally convex Four subgroups are distinguished on the basis of: (1) continuity of laminae across the zooecial boundary zone, (2) occurrence of subgranular calcite, and (3) occurrence of thick zooecial linings In the second group, laminae intersect the axis of oral growth at less than 90° In one subgroup, laminae are linear to slightly curved; in a second subgroup, laminae recurve aboraliy to form a broad arch in the outer cortex The last subgroup occurs in a Bathonien species, thus extending the known occurrence of orally acute lamination ; Bulletin 291 ACKNOWLEDGMENTS This study was undertaken as a doctoral thesis at the University of Cincinnati under the guidance of K E Caster Research was carried out at the National Museum C, under the direction Washington, D C, was made ington, D in Program of in of of R S Natural History, Wash- Boardman The research possible through the Cooperative Paleontology which exists between the National Museum Natural Histor}^ and the University of Cincinnati The author grateful for financial assistance provided is by the Smithsonian Re- search Foundation; and the grants to defray publication costs from the University of Cincinnati and W^ayne State University Sincere thanks are extended to the following: Jesse Merida, David Massey, Lorenzo Ford and tional Museum A H Cheetham (National Museum of F Donald Dean, J Collier (Na- Natural History) for discussion of techniques; of Natural History), John Pojeta, Ellis Yochelson (United States Geological Survey) for dis- problems; members of the Seminar on cussion of nomenclatural Bryozoa at the National Museum of Natural History including R S Boardman, A H Cheetham, O L Karklins, T G Gautier, R W Hinds, R J Scolaro, and R J Singh for advice and sugges- John Petering (Wayne State University) tions; gram and processing data Center; many the at the Wayne Romach (Wayne State text figures; Patricia M Nye Eileen of the manuscript; Erhard Voigt for writing a pro- State University University) Computer for drafting for typing and improving (Geologische-Palaontologisches Hamburg), Emil Buge (Museum National d'Histoire Naturelle, Paris), P A Cook (British Museum, Natural History), Horace Richards (Academy of Natural Sciences of Philadelphia), Uday Bagwe (Yorkshire Museum), Heinz Kollman (Naturhistorisches Museum, Wien), Arnfrid Durkoop (Universitat, Bonn) for loaning specimens and for allowing thin-sections to be made of Institut, critical specimens Collection of European localities was supported by a grant from the Treatise on Invertebrate Paleontology and from the Smithsonian Research Foundation I am grateful to the following individuals for their help in collecting these localities: L J Pitt England), John Neale (Hull University), (North Harrow, (Geo- Erhard Voigt Cerioporid Cyclostomes (Bryozoa): Nye Hamburg), H W, J v.Amerom (Netherlands Geological Survey, Heerlem), Emil Buge (Museum logisches-PalaontoIogisches Institut, National d'Histoire Naturelle, Paris) ABBREVIATIONS FOR REPOSITORIES USNM National States Museu mof Natural History (formerly United Museum), Smithsonian Institution, National Washington, D C BM British Museum (Natural History), London, Great Bri- tain MNHN Institute de Paleontology, Museum National d'Histoire Naturelle, Paris, France NMW Naturhistorisches UB Institut ANSP Museum, Vienna, Palaontologie, Universitat, public of Germany Academy of Austria Bonn, Federal Re- Natural Sciences, Philadelphia, Pa., U.S.A INTRODUCTION Fossil genera of Cyclostome Bryozoa have been known since 1826 when Goldfuss erected the genus Ceriopora Since that time, numerous cyclostome genera and species have been named, particularly in the works of Michelin (1841-1848), Haime (1854), von Hagenow (1851), d'Orbigny (1849b, 1854), Gregory (1896, 1902, 1909), and Canu and Bassler (1920, 1922, 1926) Knowledge of living cyclostomes has been increased by the efforts of Barrois (1877), Busk (1879), Waters (1879), Harmer (1890, 1893, 1897, 1899), Robertson (1903, 1910a, b), and Borg (1926a, 1933) The abundance of named species and genera and the length of time that they have been known suggests that cyclostome bryozoans should be, at present, a well-known group taxonomically Yet this is not the case Since the beginning of this century, cyclostomes have largely been relegated to the backwaters of taxonomic research With the exception of Borg's investigations, fundamental understanding of cyclostomes has not advanced since about the turn of this century The major obstacle to the investigation of cyclostomes has been the lack of study techniques In the past, were based on a few most taxonomic studies arbitrarily chosen external characters Taken Bulletin 291 singly or together, these characters were generally non-diagnostic virtue of: a) their ubiquity throughout the cerioporids, tubes cylindrical to prismatic", b) their ambiguity, tubes long", or c) their having so much e.g., by "zooecial "zooecial e.g., intertaxon variability as to be virtually useless Definitions of taxa were unreliable and have not served to define or distinguish taxa Illustration of the external characters of types has failed to provide sufficient documentation at homeomorphy, external characters are poor data from which to infer evolutionary relationships As a result, existing taxonomic frameworks are inconsistent and largely unuseable Thus, cyclostomes have been virtually ignored in geologic or biologic investigations which depend the specific or generic level Furthermore, largely because of upon taxonomic information This study is as basic data an attempt to find new characters that will provide the data for the construction of a new taxonomic framework One of the finest collections of fossil cyclostomes in the world is housed in the National Museum of Natural History Numerous cyclostome species were thin-sectioned under the direction of R S during the summer Boardman of 1966 Preliminary examination of these sec- tions indicated that cyclostomes have at least as many internal char- acters as Paleozoic Stenolaemata Species with relatively large or "stony" zoaria were easily thin- sectioned by techniques in general use Many of these species were and the most recent comprehensive cerioporid genera was given by Bassler (1953) Theretreatment of fore, the genera selected for this initial study were those assigned by Bassler to the Cerioporina as valid names or synonyms referable to the Cerioporina, TAXONOMIC BASIS AND PROCEDURE APPROACH The major is nomennames and goals of this revision are two-part.The first clatural: to determine the validity of generic and specific document types, primarily through photographic illustrations Types are the objective fixtures of nomenclature and must form the nucleus of any revisionary taxonomic investigation Validation of generic names was facilitated by the large collection of literature on bryozoans collected by R S Bassler, later R S Boardman and A H Cheetham, and by the large general colto Bulletin 291 216 Explanation of Plate 47 Page Figure Zonopora spiralis (Goldfuss), 1826 156 Lectotype UB 133 Specimen is probably the specimen figured by Goldfuss, 1826, pi 11, figs 2a, b, from Cretaceous, Maastrichtian, St Petersberg near Maastricht (Limburg), Netherlands Branch X5 showing The convexly curved typical helical appearance of zoaria zoarial surface is referred to as the zoarial salient; the concave surface as the zoarial embayment Terminal diaphragms appear as a nearly continuous sheet over the upper portion of the embayment lb Surface of branch X20, detail of la Ic Deep tangential thin-section X30 Id Tangential thin-section XlOO Deep tangential section of la large zooecia showing amalgamate laminate, zooecial linings le If and thin, Transverse thin-section X30 Large zooecia on lefthand side opening at zoarial salient, small zooecia on right opening at zoarial embayment Branch axis is offset to right center Longitudinal thin-section X30 Large zooecia directed towards zoarial salient on right, small zooecia towards zoarial embayment on left Note terminal diaphragm in small zooecia at Ig appearance left Tangential thin-section X30; relatively shallow section of large zooecia showing thick zooecial linings Note spinelike projections of homogeneous-appearing cortex tissue towards zooecial chamber Mural spines are commonly submerged by thick zooecial lining in outer exozone Bull Amer Paleont., Vol 69 Plate 47 I ^i^* c ;^^^*^^:^.: i.^.^^ c>v^- ^fev pi - I g Bull Amer Paleont., Vol 69 Plate 48 Cerioporid Cyclostomes (Bryozoa): Nye 217 Explanation of Plate 48 Page Figure Zonopora USNM spiralis (Goldfuss), 1826 2965A-25, Cretaceous, (Limburg), Netherlands la Loc 156 Maastrichtian, Maastricht XS Shows profile with and embayments resulting from helical major vector of distal growth Large zooecia Longitudinal section, acetate peel zoarial salients growth about a intersect surface of salient at nearly right angles; small zooecia intersect surface of erabayment obliquely Branch to lower left appears to be an intrazoarial overgrowth lb Tangential section, acetate peel X30 Shallow tangential section of small zooecia, deep tangential section of large zooecia Ic Longitudinal section, acetate peel X30, detail from la Zooecial walls in endozone thin and parallel-sided Exozonal zooecial walls thicker, sometimes submoniliform, and have characteristic lanceolate profile Id Longitudinal section, acetate peel XlOO, detail of Ic showing typical wall structure with light-colored cortex tissue bounded by thick, dark-colored deposits of zooecial lining Spinelike lateral projections of cortex tissue submerged by zooecial lining, but sometimes project into zooecial cavity as mural le If spines Longitudinal section, acetate peel XlOO showing wall structure of small zooecia and interzooidal pores Note thick deposits of dark-colored, longitudinally laminated tissue Tangential section, acetate peel XlOO Section is just distal to le, showing shallow tangential view of small zooecia Note amalgamate, granular appearance of zooecial walls and thick zooecial linings Ig Tangential section, acetate peel XlOO Section is moderately deep, showing thin zooecial lining Note mural spines and narrow, straight, interzooidal pore in lower left quadrant of figure Bulletin 291 218 Explanation of Plate 49 Page Figure Zonopora USNM 296SA-7, Cretaceous, (Limburg), Netherlands la 156 spiralis (Goldfuss), 1826 Loc Maastrlchtian, Maastricht Longitudinal section, acetate peel X30 section, acetate peel X30 Large zooecia on small zooecia on right Axis of branch is shifted to right of center; note thick zooecial linings in exozone Ic Tangential section, acetate peel XlOO, detail of large zooecia in lb Note increased thickness of zooecial lining in exozone Id Tangential section, acetate peel X30 Upper part of figure shows shallow tangential section of small zooecia of zoarial embayment and deeper tangential view of large zooecia in lb Transverse left, zoarial salient le Longitudinal section, acetate peel XlOO The zooecial boundary zone is marked by a narrow zone of light-colored, homogeneous tissue bounded by the indistinctly laminated cortex tissue Lamination in the cortex, as seen in second zooecial wall from right, is only slightly curved convex orally, and abuts the zooecial lining at 60°-70° Zooecial linings are characteristically thick and dark in color A narrow, straight, interzooidal pore is seen in aboral part of second zooecial wall from right Mural spines extend into zooecial cavity in more aboral portion of zooecial wall in left part of figure Spinelike extensions from cortex are submerged by thick zooecial linings in zooecial walls in right- hand portion of figure Bull Amer Paleont., Vol 69 Plate 49 Bull Amer Paleont., Vol 69 Plate 50 2C Cerioporid Cyclostomes (Bryozoa): Explanation of Plate Nye 219 50 Figure 1-3 Page Zonopora spiralis (Goldfuss), 1826 specimens from Cretaceous, Maastrichtian, Maastricht burg), Netherlands All 156 (Lim- USNM Loc 2965A-21 Longitudinal section, acetate peel X30 showing small zooecia at zoarial embayment with porous terminal diaphragms Primary growth is encrusted by intrazoarial overgrowth lb Longitudinal section, acetate peel XlOO, detail of la Terminal diaphragm is porous and similar in appearance to zooecial lining of subjacent zooecial walls Adjacent diaphragms are not continuous across terminus of zooecial wall Light-colored, homogeneous tissue of basal layer is continuous with cortex tissue of encrusting zooecia Loc 2965A-23 2a Longitudinal section, acetate peel X30 Shows large zooecia budding from axial region offset to right side of branch Shows thin, parallel-sided walls in endozone and thickened exozonal walls with characteristic club or lanceolate profiles Note that the apertural rims are constricted, forming a cusp-shaped profile from which terminal diaphragms extend la USNM laterally 2b Longitudinal section, acetate peel XlOO Tissue of terminal diaphragm is continuous with zooecial linings, but does not extend across light-colored cortex tissue at terminus of 2c Longitudinal section, acetate peel XlOO Detail of la showing cuspate extension of apertural rim and lateraj flexure of zooecial lining to form terminal diaphragm Loc 296SA-26, longitudinal section, acetate peel XlOO Zooecial walls of large zooecia in outer exozone The zooecial wall USNM and rodlike extension of cortex into the mural spines, suggest a younger ontogenetic stage than shown in PI 36, figs Id, e, and pi 37, thin zooecial lining zooecial cavity as fig le Bulletin 291 220 Explanation of Plate 51 Page Figure Recent cerioporid, USNM 6086-1, X30 Brood chamber with roof partially broken away to show interior, revealing intrachamber zooecia connected by thin septate walls O'Donoghue Coll 1963-2-6-1 pt all Recent cerioporid, XlOO Soft tissues are stained 2a Tangential thin-section Basal portion of tentacular crowns 38 BM in three Dark 2b 2c zooecia; lines in brown bodies skeletal wall in smaller zooecia to left probably are algal or fungal borings Note dark staining nuclei of cellular tissue in interzooidal part indicated by arrow Longitudinal thin-section Shows tentacular crowns and visceral sacs of two zooids Insertion of lophophore retractor muscles directly on body wall shown in zooid on right; note lack of funiculus at base of visceral sac Shows tentacular crown, visceral sac and brown bodies in center zooecium Nucleated strands of tissue appear to pass continuously through interzooidal pore marked by arrow 51 Plate 51 Bull Amer Paleont., Vol 69 2b 2c INDEX Note: Light face figures refer to page numbers Bold face figures refer to plate numbers A 63 Acamptostega Acanthoceramo- Ditaxia 13,87,88,93, 114 20-22 53,87,88-95, 189-191 88 88 Heteropora Polytaxia B Berenicea beyrichi, Pennipora 53, 114, 148, 150, 154 113, 147, 148, 150, 214, 215 Heteropora 30 porella anomalopora, Ceriopora dichotoma, Ceriopora Tetrocycloecia 14,147,148- 45,46 155, 214, 215 13, 23, 27, 32, 80-81, 97 140 13, 14, 21, 27, 87-88, 113, Diplocava Ditaxia Domopora 44 213 56, 61, 176- dujardini, Fungella Dysnoetopora 114 63,64 63 10, 13 178 F C 64 122 130 97 30 Calyptrostega canui, Multigalea Cavaria Cellaria Ceramoporella Ceriocava 12, 22, 27, 30, 4043, 61, 79, 81, 98, 108,139 falax, Ceriopora frustulosa, 27, 52, 56, 61, 87, 112, 113, 120, 139, 140 8, 40, 64, 70 140, 145, 147 corrugata, Densipora 131 Cerioporina conifera, Millepora Corymbopora fungiformis, Reptomulticavea Corymbosa corymbosa, Ceriocava 1-6 23, 32, 41, 42, 43-52, 67, 99, 108, 170-175 Millepora Coscinoecia Ceriopora 14,53,55, 139-141, 145 Corymbopora Diplocava Reptonodicava 41-44 60, 61, 176- Brachysoecia 33 53, 154 H Haploecia 10, 11, 13, 22, 27, 32, 33, 41, 81,96-98, 112 Heteropora 14, 21, 28, 5255, 61, 79, 87, 178 cryptopora, Ceriopora Heteropora 17, 55, 140147, 210-213 Grammascoecia 79 Heteropora 17 122 grandis, 38, 40, 43, 67, crassa, 122 G 12,40,42,43 12, 22, 27, 30, globosa, 10, 12, 27, 62- 64 12,62 42 63,64 Monticulipora Fungella 7,11,12,21, Ceriopora 122 62-64 Fasciculipora 112-115, 120, 14,112,113, 32-35 24, 53 54, 56, 67, 114, 115122, 201-204 147 64 114 Heteroporina Heterotrypa D Defrancia Dendroecia 63,64 incondita, Diplocava 13, 41, 96, 98, 112 221 16-19 13, 81-27, 122, 185-188 INDEX jacksonica, Parleiosoecia 37-40 14, 50, 67, 129-132, 135138, 206-209 lamellosa, Multicrescis Leiosoecia 122 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