Pennie wa AR niet . -— HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology 2, Py a: BOLLETINS OF AMERICAN PaRPONTOLOGY (Founded 1895) UIRRARY : FEB 17 1976 MH cAN re VA FF ES UNIVERSITY Vol. 69 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 14850, U.S.A. PALEONTOLOGICAL RESEARCH INSTITUTION 1975-1976 PRESIDE NUD esos ck coc ean en ee ene Oe Be ae Eo HArROoLp E. VOKES AV;T@E=P-RESTD EIN Ty sees re ee ee eee ae eres eI re DUANE O. LEROY SEGRE CARY coe rece ee wae ey tee ea ae ee Re ee ee ee PHILIP C. WAKELEY TDIRECTORS| (DI REASURER seco esos oo sarong oe es ocean eee KATHERINE V. W. PALMER (ASSISTANT DD TRE CO OR) oe cies ee oe a ence en ae ee Davip W. KIRTLEY ASSISTANT SECRETARY, ASSISTANT TREASURER ............-----------eeceeo+e= REBECCA S. HARRIS COGN GED hese a cee See A ee Oe ARMAND L, ADAMS IREPRESEN/ DATIVE AvAU Atm © O INC Ligne sess RICHARD G. Oscoop, JR. Trustees RutTH G. Browne (1974-1976) KATHERINE V. W. PALMER (Life) KENNETH E. CAsTER (1975-1978) JoHN PojeTA, Jr. (1975-1978) MeErRILL W. HAAs (1973-1976) CASPER RAPPENECKER (1973-1976) Resecca S. Harris (Life) K. NorMAN SACHS, JR. (1974-1977) MarcGareT B. HeEroy (1975-1978) DANIEL B. Sass (1974-1977) Davip W. KirTLEY (1974-1977) Harotp E. Vokes (1975-1978) DUANE O. LERoy (1974-1977) PuHitip C. WAKELEY (1973-1976) AXEL A. OLsson (Life) BULLETINS OF AMERICAN PALEONTOLOGY and PALAEONTOGRAPHICA AMERICANA KATHERINE V. W. PALMER, Editor Doris C. BRANN, Assistant Advisory Board KENNETH E. CASTER HANS KUGLER A. Myra KEEN Jay GLENN Marks AXEL A. OLSSON Complete titles and price list of separate available numbers may be had on application. For reprint, Vols. 1-23, Bulletins of American Paleontology see Kraus Reprint Corp., 16 East 46th St., New York, N.Y. 10017 U.S.A. For reprint, vol. I, Palaeontographica Americana see Johnson Reprint Cor- poration, 111 Fifth Ave., New York, N.Y. 10003 U.S.A. Subscription may be entered at any time by volume or year, with average price of $20.00 per volume for Bulletins. Numbers of Palaeontographica Ameri- cana invoiced per issue. Purchases in U.S.A. for professional purposes are de- ductible from income tax. For sale by Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850 U.S.A. BUIREE FINS OF AMERICAN PALEONTOLOGY (Founded 1895) Vol. 69 No. 291 A Lewis G. WEEKs PUBLICATION GENERIC REVISION AND SKELETAL MORPHOLOGY OF SOME CERIOPORID CYCLOSTOMES (BRYOZOA) By OsBorNE Barr Nye, Jr. February 12, 1976 Paleontological Research Institution Ithaca, New York 14850, U.S.A. Library of Congress Card Number: 75-44572 Printed in the United States of America Arnold Printing Corporation Ithaca, N.Y. CONTENTS Page TaN OTS Eo 0 ee Peete ARETE EB eet eee sg SEE Sit, 8 VCS a pe a A ae eee 5 PENCE Ue AK oXa Kg 11 251 pepe RP Se ee ee ee 6 AD Dre waations fOrenre PO Sit les) cc eters scorer ee ae eee eee 7 Minn te OU UAC tL O Tn ge ces et sca oss a cca aa a acai oo SEE oer pcan 7 WaxOnomicsbaSis) and! proCe Cum e mcsss eee ere ees oer eee ae eee 8 JaXfoy 0K: KG Var este cane tae eee sneer nee ee eee ROO eM ee eee Ee ee err 8 Glenera sinc hide die: Sh ere ere tie ene eae e ne en eee ee ene renee 19 Synonymies ................ Soe ne i an Pate ery ad roe A ego eae es Cee een eS ee 10 GENE TIGUGLACT OSE Sere sss a a eae es eC 11 Tie CLC GUC Seek earn er erecta ne Neen EER Ce oe rea ee eee 11 IB YOMTIE ER ICS: eee tae eee ae eee Ee BRR ere tee See eae aere ee 15 Skeletal@monphol O piyse es 2.8 occ cxtecsey me eae eer ee erry Se 18 ZOOeCI ally awWialll eS GU GEU Tey care Nee lene sree ace ete ee a Oe et ce 18 INVA C RO SEDUCE Ceres ee ee a ee ee Er 18 Orally oblique lamination ....................... Sle eee ee Bin SE a 21 Orally acute. lammaty om iscsssessr sees ae nc ee 23 Wiarrratrone ins thickmes sic cscs ecco ee ceceaesces e ee e 23 Diaphragms-and simple external) walls es. ee 24 Introduction ssc ks eek vie: ee, Pe ee ee ee 24 Inter zoordall pOnesxes ceo ean ol Oo ees, oe Re ee 35 Zoariall brOo der han be 15 ees eee ae ee aed eee 38 Systema tema ese riptro mses cee cse eee eee er tee Seca es eee Ee 49 Reference sin cites cs 2-2: bate am cee Oe No te a ee Oe el, See eel 162 DPT eat Sige wee ce ee I eS See et IE net 169 1 Las (oS oa eee Eh EPR Seni tM ab. ol ot) Set MRR rnd Om SM ewe a eNe re Seo duip SE) 4. 07/7) ILLUSTRATIONS Text-figures 1-20 TABLES Tables 1-30 ~ é Sy! aie ee wt ey ee" a stile 448 dowd ) oa? oth i — Me ° = alta “i ’ audi F ‘am, oileg re _ i” ici Ves WL 7 ) » joa é ee. y 4 ei i b=" aed e& an | 1 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 contro] 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 dia- phragms. Brood chambers, which are primary zoarial structures in the cerio- porids 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 auto- zooecia 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 intrazoarial 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 distin- guished 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 wal] 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 aborally 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. 6 BuLLeTIN 291 ACKNOWLEDGMENTS This study was undertaken as a doctoral thesis at the Uni- versity of Cincinnati under the guidance of K. E. Caster. Research was carried out at the National Museum of Natural History, Wash- ington, D. C., under the direction of R. S. Boardman. The research in Washington, D. C., was made possible through the Cooperative Program in Paleontology which exists between the National Museum of Natural History and the University of Cincinnati. The author is grateful for financial assistance provided by the Smithsonian Re- search Foundation; and the grants to defray publication costs from the University of Cincinnati and Wayne State University. Sincere thanks are extended to the following: Donald Dean, Jesse Merida, David Massey, Lorenzo Ford and F. J. Collier (Na- tional Museum of Natural History) for discussion of techniques; A. H. Cheetham (National Museum of Natural History), John Pojeta, Ellis Yochelson (United States Geological Survey) for dis- cussion of nomenclatural problems; members of the Seminar on 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- tions; John Petering (Wayne State University) for writing a pro- gram and processing data at the Wayne State University Computer Center; Eileen Romach (Wayne State University) for drafting many of the text figures; Patricia M. Nye for typing and improving the manuscript; Erhard Voigt (Geologische-Palaontologisches Institut, Hamburg), Emil Buge (Muséum 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 (Naturhis- torisches Museum, Wien), Arnfrid Durkoop (Universitat, Bonn) for loaning specimens and for allowing thin-sections to be made of 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 (North Harrow, England), John Neale (Hull University), Erhard Voigt (Geo- CERIOPORID CycLosTOMEs (Bryozoa): NYE 7 logisches-Palaontologisches Institut, Hamburg), H. W. J. v.Amerom (Netherlands Geological Survey, Heerlem), Emil Buge (Muséum National d’Histoire Naturelle, Paris). ABBREVIATIONS FOR REPOSITORIES USNM National Museu mof Natural History (formerly United States National Museum), Smithsonian Institution, Washington, D. C. BM British Museum (Natural History), London, Great Bri- tain MNHN Institute de Paléontology, Muséum National d’Histoire Naturelle, Paris, France NMW Naturhistorisches Museum, Vienna, Austria UB Institut Palaontologie, Universitat, Bonn, Federal Re- public of Germany ANSP Academy of 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, particu- larly 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 large- ly 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, most taxonomic studies were based on a few arbitrarily chosen external characters. Taken 8 BULLETIN 291 singly or together, these characters were generally non-diagnostic by virtue of: a) their ubiquity throughout the cerioporids, e.g., “zooecial tubes cylindrical to prismatic”, b) their ambiguity, e.g., “zooecial tubes long”, or c) their having so much 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 the specific or generic level. Furthermore, largely because of homeo- morphy, external characters are poor data from which to infer evolu- tionary 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 upon taxonomic information as basic data. This study is 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. Boardman during the summer 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 referable to the Cerioporina, and the most recent comprehensive treatment of cerioporid genera was given by Bassler (1953). There- fore, the genera selected for this initial study were those assigned by Bassler to the Cerioporina as valid names or synonyms. TAXONOMIC BASIS AND PROCEDURE APPROACH The major goals of this revision are two-part.The first is nomen- clatural: to determine the validity of generic and specific names and to 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 col- lection of literature on bryozoans collected by R. S. Bassler, later R. S. Boardman and A. H. Cheetham, and by the large general col- CERIOPORID CycLosToMeEs (Bryozoa): NYE 9 lections of zoological literature in the National Museum of Natural History. Objective documentation of genera was approached through the location and redescription of the primary types of type species. When authoritative evidence indicated that the primary types were destroyed or lost from known repositories, generic names’ were retained only if secondary specimens could be assigned with confidence to the type species. This was necessary because most concepts based on external characters generally do not serve to define or distinguish cerioporid taxa. In each instance where concepts were based solely on examination of secondary specimens, the reasons for their use are discussed. Internal characters are well known in Paleozoic tubular Bryozoa and provide the basis of internally consistent taxonomic concepts. It is reasonable to expect that the same approach should yield similar results when applied to the study of cyclostome bryozoans. The second goal has been to formulate generic and specific con- cepts based primarily on skeletal structures, and to interpret skeletal structures biologically. In this first stage of revision, numerous in- ternal structures were recognized. Choice of characters associated with certain structures does not imply inferences of phylogenetic importance but does expand the known phenotypic basis from which evolutionary inferences can be made. The concepts, if internally con- sistent, should provide the empirical data for second-level, more theoretical, studies, including the construction of taxonomies based on inferences of evolutionary linkage. Construction of phylogenetic classifications implies knowledge of variation in genotypes through time. Estimates of genetic varia- tion improve as nongenetic factors are excluded. In paleontology, variation in genotype is inferred from morphologic, primarily skele- tal, characters. Boardman, Cheetham, and Cook (1969) have identi- fied and discussed extragenetic elements which influence mode of growth in Bryozoa. These elements are ontogeny of zooids, astogeny, polymorphism, and microenvironment. Variation in these elements can be recognized in single colonies. Moreover, each colony is made up of numerous zooids and all zooids are virtually identical in geno- type. Thus, investigators of colonial organisms have a powerful tool for calibration of extra-genetic sources of phenetic variation. 10 BULLETIN 291 Taxonomic concepts, to be useful phylogenetically, should be based on characters which reflect genetic variability. Concepts de- veloped here are, admittedly, preliminary because only the types are adequately prepared for study. Species descriptions are based on few specimens, and all but one genus are based on the examination of type species only. None the less, these concepts are not invalid; they are simply imprecise. A great deal of information can be derived from a few, or even single, specimens. Types have special bearing on nomenclature, but no special bearing on concepts. They are simply members of a population and, in terms of that population, bear no more and no less information than any other individual. Concepts based on single specimens pose a special problem be- cause concepts nominally imply knowledge of interspecimen vari- ability. Herein, two species are presently known from lectotypes only. Because the specimens showed states of many characters as- sumed to have importance in other species, and because estimates of nongenetic variability can be made even from single zoaria, these specimens were fully described. GENERA INCLUDED Of the approximately 50 cerioporid generic names listed by Bassler (1953), 17 are listed in Table 1 and represent progress to date on the generic revision of the group. Of the 19 names, four are objective synonyms, one is a subjective synonym, and one genus, Dysnoetopora, has been reassigned to the Cheilostomata (Voigt, 1971). The 13 remaining genera show relatively great variation in mode of growth and wall structure. In the future, it may be neces- sary to remove two of them (Corymbopora and Haploecia) from the cerioporids. Reassignment is not made here because all genera are compatible with Borg’s double-wall concept and are not re- ferable to the other existing double-walled groups, the hornerids, or lichenoporids. Many genera remain to be examined. Erection of new taxa at this stage is premature and could only serve to confuse rather than clarify. SYNONYMIES The synonymies prepared here are objective in scope. They list those works which bear on the validity of names or documentation of types. Inclusion of non-objective references bears on taxonomic concepts, and in cerioporids, morphologic concepts as presently un- CERIOPORID CycLosToMEs (Bryozoa): NYE 11 derstood here must be based to a large extent on internal characters. Earlier investigators have based concepts on the relatively few ex- ternal characters. Thus, published descriptions and illustrations are not sufficient for evaluation. Relatively complete synonymies for names proposed prior to about 1900 are listed by Gregory (1896, 1899, 1909). GENERIC DIAGNOSES Generic diagnoses, excepting that for Haploecia Gregory, are based on the type species. Information concerning specimens actually examined in this study is summarized in Table 1. Characters (or character groups) believed to be useful at the generic level are: 1) Zoarial growth patterns, including the occurrence of poly- morphism. 2) Microstructure of the zooecial wall. 3) Occurrence of diaphragms, and simple external walls. In order to maintain consistency, characters based on structures observed in relatively few genera were excluded from generic diagnoses but were included in species descriptions. Brood chambers, for example, are striking morphological structures which are easy to identify and often have characteristic shapes. As such, various authors have considered them as important taxonomic characters at nearly all subordinal ranks (e.g., Canu and Bassler, 1920). In this study, brood chambers were observed in only five genera, and possibly a sixth (large primary chambers were observed in Cerio- pora Goldfuss, but other structural characteristics typical of brood chambers were not observed). In four genera, the brood chambers were abundant and many occurred in each specimen. In the re- maining genera, brood chambers were few; in Parleiosoecia Canu and Bassler, only three brood chambers were seen in 30 specimens. Brood chambers, therefore, were not included in generic diagnoses. It is hoped that future investigations will clarify the occurrence and taxonomic importance of these structures. TECHNIQUES When this investigation was begun, standard thin-section and peel techniques, as modified by R. S. 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At that time, poorly indurated fossil cyclostomes and non-indurated Recent speci- mens were vacuum impregnated with polyester resins. In the course of this investigation, modifications of these techniques were made (Nye, Dean, and Hines, 1972). Essentially, these amounted to the utilization of epoxies for impregnation and mounting, and included fine polishing procedures of cut and ground faces. These modifica- tions resulted in improved resolution of internal structures, the ability to section hard and soft parts together in Recent specimens, and the ability to make thin sections when desired (to approximately 5 microns). BIOMETRICS Numerous characters were measured. A listing of these charac- ters is given in Table 2. Phrases describing particular measurements are not always brief. Therefore, it was necessary to use abbreviations in statistical summaries of measurements found in each species description. The abbreviations are listed in Table 2. Measurements of micro-dimensions were made directly through the microscope using an ocular micrometer. Projection techniques, which are faster, were attempted initially, but had to be abandoned because projected images of many specimens lacked sufficient con- trast. Commonly, more zooecia are available for measurement in tangential sections than it was feasible to measure, so a method of selection was necessary. Non-random methods of selection intro- duce bias and place constraints upon parametric statistics. Two random methods of selection were designed and are described below: 1) The microscope stage used could be moved parallel to two directions at right angles. A scale on the stage, calibrated to .1 mm, indicated the distance in each direction. The section to be measured was positioned, and the coordinates of the corners of a four-sided polygon which enclosed most of the section were noted. These co- ordinates were transferred to graph paper and a grid was constructed. 16 BULLETIN 291 TABLE 2 KEY TO ABBREVIATIONS USED IN STATISTICAL SUMMARIES Zoarial Zr-Ht Zr-Wth Br-CsSn-MxDn PrBr-CsSn-MxDn Ov-Th AxCh-CsSn-MxDn BslLyr-Th BrCpm-MxDn BrCpm-MnDn Zooecial ZcCh-CsSn-MxDn ZcCh-CsSn-NMxDn Zc-LgnSn-Dph ZcCh-CsSn-MxDn ZcCh-CsSn-NMxDn CdZcW1-Th ZdPr-Cn/ZcCsSn ZdPr-MnDr CnlZcCh-CsSn-MxDn ZwWI1Ln-Th ZcSp-Cn/ZcCsSn ZcCh-CsSn-LgnDn ZcCh-CsSn-TrvDn Simple External Wall SEW-Th SEW-Pst-CsSn-MxDn SEW-Psdp-CsSn-MxDn Diaphragm TrlD-Th TrlD-Psdp-CsSn-MxDn IntD-Th IntD-Intv] IntD-DncApt BsID-Th Bs!ID-Intvl Zoarial Height Zoarial Width Branch — Cross Section — Maximum dimension Primary Branch — Cross Section — Maximum dimen- sion (intrazoarial) Overgrowth — Thickness Axial Chamber — Cross Section — Maximum Dimen- sion Basal Layer — Thickness Branch Capitulum — Maximum Dimension Branch Capitulum — Minimum Dimension Zooecial Chamber — Cross Section — Maximum Dimension Zooecial Chamber — Cross Section — Normal (to) Maximum Dimension Zooecium — Longitudinal Section — Depth Zooecial Chamber — Cross Section — Maximum Dimension Zooecial Chamber — Cross Section — Normal (to) Maximum Dimension Compound Zooecial Wall — Thickness Interzooidal Pore — Count/Zooecial Cross Section Interzooidal Pore — Minimum Diameter Central Zooecial Chamber — Cross Section — Maxi- mum Dimension Zooecial Wall Lining — Thickness (intra) Zooecial Spines — Count/Zooecial Cross Section Zooecial Chamber — Cross Section — Longitudinal Dimension Zooecial Chamber — Cross Section — Transverse Dimension Simple External Wall — Thickness Peristome in Simple External Wall — Cross Section — Maximum Dimension Pseudopore in Simple External Wall — Cross Section — Maximum Dimension Terminal Diaphragm — Thickness Pseudopore in Terminal Diaphragm — Cross Section — Maximum Dimension Intermediate Diaphragm — Thickness Intermediate Diaphragm — Interval Intermediate Diaphragm — Distance (from) Aperture Basal Diaphragm — Thickness Basal Diaphragm — Interval CeRIopoRID CycLostomes (Bryozoa): NYE Brood Chamber BrCh-Lth Brood Chamber — Length BrCh-Wth Brood Chamber — Width BrCh-Dth Brood Chamber — Depth BrChFI1-Th Brood Chamber Floor — Thickness BcChRf-Th Brood Chamber Roof — Thickness BrChPsdp-Dr Zoarial Position Pseudopores (in) Brood Chamber Roof — Diameter NO Not observed Ex Exozone En Endozone LgnSn Longitudinal Section TngSn Tangential Section TrvSn Transverse Section Statistics OR Observed Range xX Mean S Standard Deviation CV Coefficient (of) Variation N Number (of observations) NZc Number (of) Zooecia NZr Number (of) Zoaria 17 Each locus on the grid had an x and y coordinate. The number of zooecia to be measured was selected; then coordinates were chosen from a table of random numbers. The slide was positioned with re- spect to these coordinates, and the zooecium nearest the center of the field was measured. This method was time-consuming, as each tangential section required the construction of a new grid. Also, if zooecia were small and the zooecial wall thick, the coordinates were imprecise. This method was used to select zooecial characters in Reptonodicava globosa (Michelin) but was abandoned later in favor of the second method. 2) A photograph of the section was made and zooecia were num- bered directly onto the photograph. Then numbers were selected from a table of random numbers. The zooecia so chosen were measured directly through the microscope. This method is fast; a polaroid 4x5 camera back was used, and prints were available within seconds. The method is precise, as well; if measurements are suspect, the zooecium can be found and dimensions checked. Some measurements of zooecial characters are illustrated graphically for each species except Corymbopora menardi Michelin. The dimension normal to the longest dimension of the zooecial chamber was chosen because it should not be influenced by the 18 BULLETIN 291 angular relation between the plane of the section and the zooecial growth axis. Also included are histograms of the ratio of major zooecial dimensions, compound zooecial wall thickness, and a cumu- lative curve for interzooidal pore counts. Estimates of the arithmetic mean (X), the standard deviation (S), and coefficient of variation (CV) are not given for counts of interzooidal pores per zooecial cross-section. These counts do not meet the basic assumptions required for the use of parametric sta- tistics; most importantly, when plotted, they do not approximate a normal distribution. The counts are summarized in cumulative curves given for each species. SKELETAL MORPHOLOGY ZOOECIAL WALL STRUCTURE MICROSTRUCTURE Since the major studies by Ulrich commencing in the 1880's, wall structure has been considered an important taxonomic character in studies of Paleozoic stenolaemates. Nicholson was probably the first to make oriented thin-sections and observe skeletal microstruc- tures in cyclostomes. He recognized and figured the laminar structure in the zooecial wall of Recent cerioporids (1880, p. 335; text-fig. 2, p. 336), Bleicher (1894, pp. 99-100, pl. 1, figs. 1, 3; pl. 2) prepared thin-sections and illustrated laminar structure in the zooecial wall of an encrusting tubuliporid cyclostome. Later investigators have misunderstood, or virtually ignored, microstructure. In cerioporids, calcareous zooecial walls between adjacent zooecia are compound because they are grown from both sides. Therefore, in most genera, zooecial boundaries cannot be precisely defined because they lie within broad, tangentially-amalgamate zones. Laminae are sometimes arched continuously across the zone, or the zone may be composed of light-colored, subgranular, skeletal material which is nearly homogeneous in appearance. The continuity of calcareous tissue across the zooecial boundary zone suggests that the depositing epithelium passed continuously over the rims of ad- jacent zooecia. A membrane that included an outer cuticle covers the zoarium (observed by Borg, and probably Waters and Busk in Recent cerioporids, and by Harmer in Recent lichenoporids). The CrerIoporip CycLosTomMEs (Bryozoa): NY& 19 outer membrane (gymnocyst of Borg) protects the inner depositing epithelium and probably aids in the transfer of nutrients around the actively growing apertural rims. Narrow, well-defined zooecial boundary zones can be seen in only a few genera. In these genera, the laminae of adjacent zooecia meet at relatively low angles. Wall structure is not homogeneous throughout a zooecium. Zooecial walls are commonly homogeneous to subgranular, sometimes vaguely laminate in the thin-walled endozone and inner exozone portions. Thin zooecial linings are commonly present throughout. These are generally composed of dark-colored, longitudinally parallel laminae. Zooecia which bud from basal layers often have thick zooecial linings at the proximal tip of the zooecium and along the recumbent zooecial wall (PI. 39, fig. 5). Borg illustrated linear structures in the calcareous walls of Recent cerioporid’ species (1933, text-tigs. 11, 15, 16, 17; pl. 7, figs. 5, 6). He referred to these, however, as fibers (1933, p. 337) and believed that they were organic, unspecified (¢.g., see 1926a, p. 196), or chitinous (1926b, p. 585). Recently, an integrative model of zooecial wall growth in Steno- laemata was presented by Boardman and Towe (1966, p. 20). The model was more fully developed by Boardman and Cheetham (1969, p. 211, text-fig. 2, p. 210). A similar approach has been used by Tavener-Smith (1969) and by Brood (1970a). This model inte- grates Borg’s observations of the membranous portions of the body wall and Boardman and Towe’s observations of microstructure and ultrastructures of the calcareous wall. Three-dimensional laminar configuration is the principal key to the understanding of skeletal morphology. Provided that the primary lamination is preserved, one can interpret structural relationships, sequence of events, and the location of the depositing epithelium (Boardman and Cheetham, 1969, p. 210). This model provides the basis for an understanding of the growth of zooecial walls in cerioporid bryozoans. In the outer exozone of cerioporid genera, two major kinds of laminar microstructure can be distinguished. In one group, laminae arch orally convex, intersecting the orally directed axis of growth at 90° or more (Text-fig. 1 A-D), and are orally oblique, (Boardman and Cheetham, 1969, p. 211). In well-preserved specimens, laminae are continuous across the zooecial boundary zones (Text-fig. 1 A, B) 20 BULLETIN 291 (-~S Text-figure 1 A-F. Diagrammatic profiles of compound zooecial walls in the outer exozone portion of cerioporid cyclostomes. Solid lines with arrows indicate inferred position of depositing portion of inner membrane responsible for last episode of cortex growth. Dashed lines in cortex indicate indistinct lamination; solid lines in cortex indicate distinct lamination; cross-hatching indicates subgranular to homogeneous calcareous tissue. In A-D, growth sur- faces parallel lamination, and each Jamina probably represents a single growth episode. In E and F, the growth surface parallels the depositing epithelia, but cuts across lamination; laminae probably grew by edgewise growth. Zooecial linings are included only in D. Linings may be deposited as sheetlike incre- ments, or by edgewise growth. CERIOPORID CycLostomeEs (Bryozoa): NYE yA or merge indistinctly with granular or homogeneous calcite in the outer cortex (Text-fig. 1 C, D). Each lamina is inferred to have been a simple growth surface which paralleled the depositing epithelium (Boardman and Cheetham, 1969, text-fig. 2A, p. 210). In the second group, laminae intersect the axis of oral growth at less than 90° (Text-fig. 1 E, F) and are orally acute. Laminae meet with an angular relationship along the zooecial boundary zone producing an integrate appearance in tangential section (PI. 18, fig. 3). Boardman and Cheetham (1969, p. 211, text-fig. 2B, p. 210) showed that laminae in this configuration were not parallel to the depositing epithelium and thus do not constitute single-event growth surfaces. Rather, growth is simultaneous along many laminae by deposition of calcareous crystals on the leading edge of each lamina- tion. Laminae within cerioporid zooecial walls extend aborally for only short distances and are not continuous with diaphragms. Most depositional activity, therefore, takes place at, and near, the apertural rim, and the deposition of diaphragms cannot be correlated with depositional events in the compound zooecial wall. Conversely, the zooecial walls of many trepostomes are composed of laminae which can be traced long distances aborally from the aperture. Often these laminae are continuous structurally with diaphragms, and form sin- gle diaphragm-wall units (Boardman, 1969, p. 27, text-fig. 8, text- fig. 10, p. 31). In these, the membrane lining the entire living cham- ber apparently acted as a single depositional unit. In cerioporids, microstructural subgroups can be distinguished. These are described below. Orally oblique lamination. — Type 1. Laminae are broadly curved, arching continuously across the zooecial boundary zone (Text-fig. 1A). In tangential view, the zooecial walls are broadly amalgamate. This pattern has been observed in Ceriopora (PI. 8, fig. 1d; Pl. 9, fig. la) Heteropora (PI. 35, fig. 2a) and Leiosoecia; and in the walls between adjacent small polymorphs in Ditaxia and Parleiosoecia (P1. 40, fig. 1f). Type 2. Laminae are broadly curved, arching continuously across the zooecial boundary zone. Laminated calcite alternates longi- tudinally with light-colored, heterogeneous to homogeneous calcite bo i) BULLETIN 291 (Text-fig. 1B). A better understanding of the light-colored calcite will necessitate investigation by electron microscopy. This tissue is inferred to be primary because it parallels well-preserved, laminated structures. The light-colored calcite forms longitudinally discontinuous plug- like bodies in Coscinoecia (Pl. 14, fig. 1f) which are lapped by laminated calcite giving an acanthopore-like appearance in tangential section (Pl. 14, figs. lg, h). These are not presently interpreted as acanthopores because the bodies are longitudinally discontinuous and because they lack structures typical of acanthopores in Paleozoic stenolaemates. Type 2 is intergradational to some extent with Type 1, but is most clearly distinguished in Coscinoecia (Text-fig. 1B, Pl. 14, fig. Le: Pl..15, fie. te): Type 3. Laminae are nearly linear in profile (Text-fig. 10, Pl. 6, figs. 1, 3a) or arched (PI. 5, figs. 1b, 1c). Laminae are distinct and closely spaced in the inner cortex. The outer cortex is light- colored and homogeneous to indistinctly laminate (PI. 5, figs. 1a, b, e; Pl. 6, figs. 1, 3a, 3b); laminae are sometimes seen to arch continu- ously across the zooecial boundary zone (PI. 5, fig. 1b). The poorly laminated appearance of the outer cortex is not simply an optical effect resulting from the angle of intersection between laminae and the plane of the section, because it was observed in longitudinal, transverse and tangential views. In addition, it was consistently ob- served in well-preserved specimens. Therefore, the appearance probably reflects some primary, but presently unknown, ultrastruc- ture. This microstructure was observed in Certocava. Type 4. The cortex is composed of light-colored subgranular to indistinctly laminated calcite. Laminae sometimes arch continu- ously across the zooecial boundary zone. In addition, the wall has a thick zooecial lining composed of dense, dark-colored, longitudinal- ly directed, parallel to wavy laminae. The lining apparently thickens through ontogeny, and smooths over irregularities on the zooecial wall, such as spinose projections. This structure was observed in Haploecia and Zonopora and is illustrated in Text-fig. 1D, Pl. 24, fig. 1f Pl. 26s hie. ld: Pl, 30; tress la, bs Piesde ue 2b, lela ete Ip; Pl. 48, fies: Id, e, 1; Pl 49) ties. tc, e; Pl. 50) tasslb, bse CrRtoporID CycLosToMEs (Bryozoa): NYE 13) Orally acute lamination. — Type 5. In profile, laminae are linear to slightly curved, and commonly intersect the orally directed zooecial growth axis at about 45° or less (Text-fig. 1E). Zooecial walls commonly are tangentially and longitudinally integrate in appearance. Thin zooecial linings composed of dark-colored, longitudinally directed laminae are commonly present. This microstructure was first illustrated by Borg (1933, text-fig. 11, p. 303; text-fig. 15, p. 322; text-fie. 16, p: (3233; text-fig, 17, p. 329; text-fig. 20, p. 3395) pl, figs. 6, 7). Crystalline ultrastructures and mode of growth of this type were discussed by Boardman and Towe (1966, p. 20) and later, more fully, by Boardman and Cheetham (1969, p. 211; text-fig. 2B, p: 210% pl. 27, figs:'fa; ib). Type 6. Laminae initially extend from the zooecial growth axis at about 45°, then are broadly arched in the outer cortex, Zooecial walls are tangentially integrate in appearance. Zooecial linings are present or absent. This microstructure was observed in Diplocava and Reptonodicava and is illustrated in Text-fig. 1F; PI. Noy ties) live heh Wiecties6bs Pl 18x fies; 2,3; PL 19st. eal 41, fig. Lf. VARIATION IN THICKNESS Zooecial walls of cerioporid bryozoans commonly show pro- nounced variation in thickness. In some genera this variation is cyclic, giving rise to annular thickenings (PI. 4, fig. le). More com- monly, however, variation in thickness shows less regular patterns. Text-fig. 2 A-D illustrate the development of several different pro- files in the outer exozonal zooecial walls of Coscinoecia radiata Canu and Lecointre. Moniliform profiles are enhanced by the occurrence of interzooidal pores but are not solely responsible for them. Inter- zooidal pores are nearly always located in thin-walled zones, but thin-walled zones are not always pierced by interzooidal pores. A quantitative estimate of variation can be made from measure- ments of the width of compound zooecial walls in tangential section. The coefficients of variation generated from these measurements range from 27 in Ceriocava corymbosa (Lamouroux) to 55 for all polymorphs in Coscinoecia radiata Canu and Lecointre. The thick- ness of individual zooecial walls could not generally be measured be- cause zooecial boundaries are not visible in thin-section. Further- 24 BULLETIN 291 more, zooecial boundaries can be roughly approximated in longi- tudinal section by noting the orally directed crest of individual laminae (Text-fig. 2). Longitudinal lines connecting these points, the zooecial growth axes, are often significantly offset from the mid- dle of the wall (Text-fig. 2C, D). Therefore, approximation of the zooecial wall thickness by halving the thickness of compound zooecial walls would often be inaccurate. Qualitative estimates of variation in zooecial wall thickness can be made from profile views (Text-fig. 2 A-D). Species tend to show characteristic profiles and apparently have certain limits to vari- ability. For instance, zooecial walls in Heteropora cryptopora (Gold- fuss) are nearly parallel-sided (PI. 34, fig. la) and become thicker orally in a regular manner. This contrasts with zooecial walls in Coscinoecia radiata Canu and Lecointre which commonly show great variation in thickness longitudinally (Text-fig. 2A, Pl. 14, fig. ia): There is some indication that this variation may be useful for delineation at supraspecific taxonomic ranks. For example, the zooecial walls near the branch tips of both Hapfloecia straminea (Phillips) and H. multilamellosa (Canu and Bassler) have irregular moniliform profiles. These become progressively more parallel-sided through the deposition of the zooecial lining. The sample, however, appears too small to generalize from, and the important elements of profile shape, e.g., its relation to micro- structure, are presently unknown. Therefore, estimates of profile shape are included here in species descriptions. DIAPHRAGMS AND SIMPLE EXTERNAL WALLS INTRODUCTION Diaphragms are intrazooecial, calcareous partitions which ex- tend transversely across zooecial tubes. They are found in the Paleo- zoic orders Trepostomata, Cryptostomata, and Cystoporata as well as the post-Paleozoic Cyclostomata. Haime (1854) was apparently the first to recognize diaphragms in the zooecial interiors of cyclo- stomes. Simple external walls, presently known only in cyclostomes, are discussed with diaphragms because they are much like terminal diaphragms in structure and position. CrrIoporip CycLtostomMes (Bryozoa): NYE 25 DISTAL GROWTH OF B ZOARIUM Text-figure 2 A-D. Incremental growth and longitudinal profiles of zooecial walls in the outer exozone of Coscinoecia radiata Canu and Lecointre. Solid lines indicate distinct laminae; broken lines indicate indistinct laminae; cross-checked patterns indicate subgranular to homogeneous-appearing tissue. Arrows indicate local zooecial growth directions. A. Profile is nearly sym- metrical across the zooecial boundary zone, but shows large variation in thick- ness longitudinally. B. Profile is nearly symmetrical across the zooecial boundary zone and is almost parallel-sided. C. Profile is moderately variable in thickness longitudinally, and subsymmetrical across the zooecial boundary zone. Note longitudinal variation in direction of zooecial growth. D. Profile is subsym- metrical across zooecial boundary zone; note longitudinal variation in direc- tion of growth. 26 BULLETIN 291 Diaphragms in cerioporids generally are constructed of super- posed calcareous laminae which are similar in appearance to the laminae of the zooecial lining. At the juncture with the zooecial wall, the diaphragms either flex orally or aborally. The orientation of this flexure can be used to infer the position of the depositing mem- brane. The flexure and depositing membrane are necessarily on the same side of the diaphragm if diaphragm laminae were deposited sequentially. In cerioporids, the flexed calcareous layers sometimes merge continuously with the zooecial lining. More commonly, the laminae adjoin the calcareous wall and extend for varying distances without merging, forming a structure referred to here as an abut- ment. Three types of diaphragms can be identified in cerioporids: basal, intermediate, and terminal. Brief descriptions of these were given previously (Nye, 1970). The occurrence of diaphragms in 13 cerioporids is tabulated in Table 3. The diaphragms can be identified on the basis of: 1) the manner in which the diaphragms join the zooecial wall, 2) position with respect to the skeletal aperture, 3) presence or absence of pores. Basal diaphragms are non-porous and generally thin, and with a few exceptions [e.g., in Haploecia straminea (Phillips), Plate 26, figure 1b], do not occur close to skeletal apertures. Basal diaphragms are primarily distinguished by the oral flexure of the diaphragm at the juncture with the zooecial wall (Pl. 5, fig. 2; Pl. 15, figs. 1b, f, 1; Pl. 43, fig. 1). This oral flexure requires that the depositing epi- thelium was oral to the diaphragms (Text-fig. 3). Basal diaphragms of cerioporids and diaphragms of trepostomes are similar because they both flex orally at the juncture with the zooecial wall. In cerioporids, orally flexed calcareous tissue commonly forms thin abutments or merges obscurely with the zooecial lining. Thus, be- cause basal diaphragms lack continuity with laminae in the cortex of the zooecial wall, deposition of basal diaphragms cannot be cor- related with depositional events in the zooecial wall. This differs from some trepostomes in which Boardman (1960, p- 27, text-fig. 8) demonstrated that, “. ..a diaphragm plus the distally connected wall deposit are interpreted to form a unit of skeletal growth that was deposited at approximately the same time . . .” Zi NYE Cer1oporip CycLostTomes (Bryozoa) usuitoads ajsuls eB UO paseg,. *SUJMOISIIAO }SOW-I9jno0 Io sdy youeiq pajsnisuaun UI B19900Z Jdadx9 UOUIWIOZ PadArasqo JON uOoUWIWIODU /) uoUIWOSU () v1920/d0 PedAtasqo JON PaArasqo JON au0zoxa !UOWUWIOOU () PpaArasqo 30N DIXDIL SYIMOISIIAO dInjiade o} 0} Jusdef[qns UCUIWIO,) PaArasqo JON esojo :uOWWOoUL:) UOUIWIOOU /) vavI01¢1q YIMOIBIIAO [BIIEOZ PedArtasqo JON PaArasqo JON -Bijul — uouwodg ‘opuq - snojauinyy 40199 0U1950/) PadArasqo JON paadrasqo 10N PaArasqo JON ‘opugq - uowo0du/) piogoqgutso;) "YJMOIBIIAO 0} JusoRfqns PedAtasqo JON Pedrasqo JON jsnf !uowwoduy pedAtasqo JON 40400149) Paatasqo JON uOWIWOD PaArasqo JON sno1auInN DVIOLI/) S[[BM [BUIa}xa a[dwig [eurWia 7 d}eIPIulsa}uy jeseg snuayy VadNAS dldOdOlddO AWOS NI STIVM IVNUALXA ATMWIS GNV SWOVUHdVIC AO AONTANNIIO £ ATaVL BuL.eTIN 291 uawtdads ajsuls eB UO pasegy 28 PeArasqo JON uOWIWO,) PeArasqo JON PeAtasqo JON vsioq¢ouoz PedArasqo JON PpeArasqo JON PeArasqo JON PaAtasqo JON D1I90]IKI04J9 T yWMoi3 I9AO 0} JUaDeEL PaArasqo ON -qns — uowUooU/) padarasqo JON sno1awinN vavripouo0jday PadArasqo JON UWOUIUIODU () uOoWW0D UOUIUIONU /) 012905019 ]40q aUu0ZO0Xxa PaArasqo JON padrasqo JON Jajno — uowwoou sy) PaArasqo JON 40199050197 asIM1ayj}O UOUWO UN SYJMOIBIIAO 0} }U99 PaArasqo JON PpedAtasqo JON -efqns — uowwo) UOUIWIOOU /) p10g041919 S][@Aa [eusa}xa a[duig [BulWia 7, 9} BIPIU19}UT [ese snuayy VUANAD GIWOdOINAO AWOS NI STIVM TIVNUALXY ATAWIS GNV SNOVAHdVIC AO AONAAANOOO CERIOPORID CycLosTOoMEs (Bryozoa): NYE 29 DISTAL GROWTH OF ZOARIUM Text-figure 3. Diagrammatic profiles of terminal, intermediate and basal diaphragms in cerioporid bryozoans. All diaphragms were deposited from the inner membrane (= cryptocyst of Borg). The inner membrane is extended over the exterior side of the terminal diaphragms; this interpretation is con- sistent with Borg’s observations of soft tissues (1933, text-fig. 2, p. 369). Arrows are emplaced at the approximate position of membranes which deposited the diaphragms. 30 BULLETIN 291 Coscinoecia has a well-developed coaxial mode of growth. Basal diaphragms are numerous and closely spaced in the endozone and the zone of zooecial bending, but they were not observed in the exozone (PI. 13, figs. 1d, g). This contrasts with most trepostomes in which diaphragms are commonly absent or infrequent in the endozone but numerous in the exozone (Boardman, 1960, p. 22). Boardman (1960, p. 34) demonstrated that “. . . diaphragm counts and width of ephebic zones can be proportional to growth stages of zooecia considering the manner of skeletal growth.” In the exozones of Ceriocava (PI. 1, fig. lg) and in Reptonodicava (PI. 44, fig. 2c), basal diaphragms are numerous and closely spaced. Ceriocava is robustly branching with ramose growth habit, and diaphragms are more numerous distally from growing tips. Reptono- dicava has a massive growth habit in which zooecia essentially grow semiradially. Zooecia growing parallel to the major axis of distal growth are long, and have numerous diaphragms; zooecia growing at an angle to the major axis of zoarial growth are shorter and have fewer diaphragms. In Reptonodicava and presumably other bryo- zoans with similar growth habits, the number of diaphragms is both a function of ontogeny and a function of zooecial growth direction relative to the major axis of zoarial growth. Intermediate diaphragms flex aborally at the juncture with the zooecial wall (PI. 9, figs. 1a, b; PI. 15, fig. 1a; Pl. 36, fig. 1g) and are the same in this regard as most terminal diaphragms. Inter- mediate diaphragms differ from terminal diaphragms in being non- porous. In addition, they are commonly thinner and are seldom ob- served at the skeletal aperture. Utgaard (1968b, pp. 1445-46, pl. 181, fig. 6) reported similar diaphragms in the ceramoporoid genera Ceramoporella Ulrich and Acanthoceramoporella Utgaard. Utgaard speculated that the diaphragms “. . . may be associated with a terminal phase of a zooid.” As noted by Utgaard (1968b, p. 1445), the aboral flexure of intermediate diaphragms requires that the soft tissues which de- posited the laminae lay on the aboral side of the diaphragm (Text- fig. 3). Soft tissues engaged in metabolic activities such as the deposition of calcareous tissue presumably require a supply of nutri- tive and respiratory substances. This, in turn, requires either a storage facility or a direct communication with tissues able to supply CERIOPORID CycLosToMEs (Bryozoa): NYE 31 these requirements. When the first calcareous lamina of the dia- phragm is completed, a chamber is formed which is sealed off from the overlying zooecial cavity. Because of the lack of pores through the diaphragm, any significant transfer of metabolites would be eliminated. In cerioporids, however, soft tissues within these cham- bers presumably have access to nutrients via the zoarial communica- tion system of interzooidal pores. Terminal diaphragms (Borg, 1933, p. 290) are deposited at, or close to, the distal extremity of a single zooecium as zooecial cover plates. They are characterized by their position: aboral flexure, forming abutments (PI. 5, fig. la; Pl. 6, fig. 3a; Pl. 40, fig. 1f) or merging with the zooecial lining (PI. 50, figs. 2b, c); and by the occurrence of pseudopores. Terminal diaphragms probably occur in most cyclostomes; they have been observed by several authors, in- cluding Busk (1859), Nicholson (1880), Waters (1884a), Pergens (1890), Robertson (1910), Canu and Bassler (1922, 1926), and Borg (1933, 1944). The epithelial tissue which deposited the terminal diaphragms was on the aboral side of the diaphragms. This is so because dia- phragms flex aborally at the juncture with the zooccial wall, and because laminae form U-shaped figures between pseudopores. This interpretation of growths leads to the inference that dia- phragms and pores were externally sealed by a cuticular layer. If so, the pores are pseudopores in the original sense of Borg and they probably functioned as cites of gas exchange. In this respect, Borg (1944, pp. 76, 116) referred to the pores in terminal diaphragms as pseudopores. This contradicts his earlier opinions in two ways: (1) he believed (1933, pp. 277-8, 289, 356) that calcite was deposited on both sides of terminal diaphragms and (2) he believed (1933, p. 368, text-fig. 26, p. 369) that ectodermal and mesodermal tissue of the inner wall (cryptocyst) lined both sides of the diaphragm. It follows that Borg must have regarded any diaphragmal pores as true pores in his original sense until his 1944 opinion. It is unfortunate that no new anatomical evidence was cited in 1944 when he called the pores, pseudopores. Terminal diaphragms are zooecial cover plates and probably serve to protect the soft tissues of the zoarial interior. In post- Paleozoic cyclostomes, the breaching of a single zooecium can pro- 32 BULLETIN 291 vide access to the entire zoarial interior via the interzooidal pores. Zoarial protection was first ascribed to terminal diaphragms by Waters (1884a, p. 403): “Now if each zooecium during its poly- pideless condition could be choked up by the mud deposited from the sea, then the whole colony might suffer”. Harmer (1890) ob- served a damaged zooecium which was sealed by a terminal dia- phragm. Later, a new polypide grew in the interior and the dia- phragm was resorbed. Borg (1944, p. 116) also observed zooecia which were broken and then sealed by terminal diaphragms. Zooids lacking feeding structures are often sealed by terminal diaphragms. Harmer (1890) observed that a “. . . zooecium which possesses a diaphragm contains a brown body, but no functional polypide. Here and there it will be noticed that a polypide-bud is being developed below the diaphragm. With the further develop- ment of this bud, the diaphragm is absorbed, the mouth of the zooecium again growing out into a long tube.” In some dimorphic species, small polymorphs which probably did not house feeding zooids are commonly sealed by terminal diaphragms (Borg, 1933, pp. 289, 368). Large polymorphs are generally open, and closure probably follows the degeneration of a feeding zooid. Perhaps external en- vironmental stresses, ¢.g., seasonal variations, may induce closure. In this case, nearly all zooecia in the zoarium would probably be sealed, as is the case observed in some zoaria of Certocava corym- bosa Lamouroux (PI. 1, figs. 1d, f). Closure would provide some protection to housed soft tissues. Presumably these tissues could survive periods of low nutrition until conditions improved, at which time feeding structures could be regenerated and the protective cover plates resorbed. Simple external walls are apertural closures which are charac- terized by the presence of pseudopores. In Haploecia Gregory and Diplocava Canu and Bassler they are, in addition to pseudopores, pierced by a single, large peristome (PI. 16, fig. 1g; Pl. 18, figs. 1-3, Pl. 28, fig. 1c). The peristome is the restricted skeletal aperture through which tentacles were probably protruded. The peristome often extends orally from the surface of the diaphragm (PL (24; fiewlt). soathe simple external walls in Haploecia Gregory and Diplocava Canu and Bassler may be homologous to the structure in the Eleidae CerroporIp CycLosToMEs (Bryozoa): NYE 33 called an “opercule” by D’Orbigny (1854, p. 606), or to the zooecial cover plates discussed by Levinsen (1912) in the eleid genus Mel- cerities d’Orbigny. The structures in Meliceritites, called opercula by Levinsen, are probably terminal diaphragms. Viskova (1968, text-fig. 16, p. 176) figured a thick section of a zooecial cover plate in Brachysoecia grandis Viskova which is probably a simple external wall. Canu and Bassler (1922, 1926) referred to the simple external wall as “facettes perforated by an aperture”. In Haploecia Gregory and Diplocava Canu and Bassler the structure of simple exterior walls resembles terminal diaphragms in that laminae commonly flex aborally at junctures with the zooecial walls to form abutments or merge with zooecial lining, and laminae form U-shaped figures between pores (PI. 18, fig. 1; Pl. 26, fig. 1a). The wall is simple because it was grown from the interior side only and the skeletal evidence requires a depositing epithelium interior to the diaphragm. Then, as argued for terminal diaphragms, the exterior side of the diaphragm was lined by cuticle and the pores are pseudopores probably functioning as cites of gas exchange. Simple external walls differ from terminal diaphragms in having restricted apertures commonly extended orally by peristomes. In ad- dition, the emplacement of the simple wall marks the end of oral growth of compound zooecial walls and are thus considered as ex- ternal calcareous walls. In many tubular bryozoans characterized by coaxial endozone-exozone mode of growth, zooecia continue to grow orally from the zone of flexure for some distance, and com- monly intersect the zoarial surface at 90°. Boardman (1960, p. 34) showed that exozones characteristically become thinner in a gradient projected distally along a branch. Boardman used this relationship as an indicator of ontogeny in the interpretation of zoarial growth. The only apparent limit to the oral growth of an individual zooectum was the death of the depositing tissue (Boardman, 1960, p. 39). In Haploecia, exozones do not show consistent change in thick- ness, apparently reaching their maximum thickness just proximal to the growing tip of the branch. Furthermore, zooecia never grow orally beyond the zone of zooecial flexure, and consequently intersect the zoarial surface at 40° to 60°, never growing beyond a zone re- ferable to the innermost exozone of many other species. Examination of unencrusted growing tips (PI. 23, figs. Ic, d) indicates that ex- 34 BuLLETIN 291 Text-figure 4 A-E. Diagrammatic profile showing inferred development of simple external walls. Shaded areas are compound zooecial wall and simple external wall; parallel lines indicate zooecial lining and peristome wall; cross-hatched pattern is basal layer and zooecial walls of intrazoarial over- growth; dark line labeled OM is outer, cuticle-lined membrane; dashed line is inner membrane; arrows indicate sites of active calcareous deposition by inner membrane; and ? indicates questionable extent of inner membrane. CEeR1opoRID CycLosToMEs (Bryozoa): NYE 35 ternal simple walls are emplaced coincidentally with the termina- tion of calcification at the oral tips of the zooecial walls. Boardman, Cheetham, and Cook (1969, explanation of text- figure 6) noted that continued growth of exozonal zooecia in a primary branch is sometimes arrested by superposition of an over- growth, and that the full ontogenetic development of zooecia in the primary branch is never achieved. The thin exozones of Haploecta straminea might be considered to exemplify the above, if only those branches with extensive encrustations were observed. Several un- encrusted branches of the type species, however, were sectioned and these also show no increase in diameter proximally. There is evidence for the continued inhabitation of the zooecial cavity by soft tissues, and continued ontogenetic development after emplacement of the simple external wall and termination of orally- directed growth of the zooecial wall. The walls are thicker, and more often structurally complex in a gradient oriented proximally from the growing tip (Text-fig. 12). Also, zooecial linings are seen to be thicker in more proximally situated zooecia. Continued deposi- tion of skeletal tissue requires the presence of depositing epidermis. In addition, peristomes probably were large enough to allow the protrusion of tentacles. It would seem possible that most of the zooecial chambers having access to the zoarial surface housed living, functional zooids. INTERZOOIDAL PORES The term interzooidal pore is equivalent to the terms inter- zooecial pore, mural pore, communication pore, infundibular pore, septulae, and canaliculi as used by other authors. Interzooidal pores are canals through the calcareous compound walls between adjacent zooids which connect the body cavities of adjacent zooids. This usage is slightly modified from the definition given by Borg (192G6a; p. 201). Interzooidal pores are not equivalent in morphology or function to pseudopores. Pseudopores pass through calcareous single walls and are bounded externally by cuticle (Borg, 1926a, text-fig. 2, p. 193). Functionally, pseudopores are thought to allow restricted communi- cation (i.¢., of dissolved gases) to the exterior, whereas interzooidal pores allow communication between adjacent zooids. 36 BULLETIN 291 The nearly ubiquitous presence of interzooidal pores in cyclo- stomes and their probable function was well known to several authors in the latter part of the last century. In 1879, Busk and Waters published separate observations on Recent cerioporid species in which they commented upon interzooidal pores. Busk (1879, p. 725) believed that the pores allowed “. . . the permeation of fluids through- out the entire zoarium”. Nicholson (1880, p. 332) noted the im- portance of the findings of Busk and Waters, and (p. 421) com- pared the structure of the pores to mural pores in the Favositidae, noting that the interzooidal pores in cerioporids have “ . . . definite walls and dilated extremities instead of being mere circumscribed deficiencies in the wall”. Examination of the microstructure of the zooecial walls in cyclostomes shows that the laminae lining the wall generally parallel the zooecial cavity until they reach the locus of the pore. Here the laminae do not stop abruptly, but deflect and contour the outline of the pore. Deflection of the laminae indicates that calcareous tissue was deposited by a secretory membrane which lined the pore; there- fore, the pore is considered a primary structure. Most Paleozoic bryozoans have nonporous walls. Interzooidal communication is limited to connections through zoarial tissues out- side of the apertures (Boardman and Cheetham, 1969, p. 214). Ordo- vician and Silurian ceramoporids have communication pores (Utgaard, 1968a, b, 1969), but they do not appear to relate struc- turally to the interzooidal pores of post-Paleozoic cyclostomes. Most post-Paleozoic bryozoans have porous walls. Waters (1884b, pp. 676- 7) believed that interzooidal pores were complete homologues of the simplest of rosette plates among the cheilostomes. The constric- tion observed by Waters, however, is not a separate structure (PI. 5, figs. le, 3; Pl. 6, figs. 2, 3b; Pl..30,-fig. 3; Pl, 51, figs. 2a-c)peand interzooidal pores differ anatomically from the interzooidal com- munications in the Gymnolaemata (Borg, 1926a; Silén, 1944; Banta, 1969). The existence of soft tissues lining the pores was first noted by Busk (1879, p. 725), and later confirmed by Harmer (1896) and Borg (1926a, pp. 201-202). Most authors believed that the inter- zooidal communication was open. Borg (1926a, p. 201), for example, wrote that, “. . . the zoids [sic] in the Cyclostomata have a much CERIOPORID CycLosToMEs (Bryozoa): NYE 37 more open communication with each other than is the case in the Cheilostomata and the Ctenostomata”. Harmer (1893, p. 213), how- ever, observed strands of funicular tissue which passed through the pores. Protrusion of the polypide is affected through increase in turgor pressure acting upon the membranous sac (Borg, 1923, pp. 7-8). If the interzooidal pores are open, increase in turgor pressure should be transmitted throughout the zoarium. Clark (1964, p. 104) con- sidered this problem and stated that, “The existence of restraints to polypide eversions is important since they permit the independent eversion and retraction of polypides which share a common hydro- static skeleton”. The restraints listed by Clark included the polypide retractors, relaxation of the vestibule dilator muscles, and contrac- tion of the vestibular sphincter. Preliminary examination of sections with both soft and hard tissues indicates that at least some pores are nearly completely filled by a single large cell (PI. 51, figs. 2b, c). Sometimes the soft tissue appears to be connected with a tenuous network of connective tissue (Pl. 51, fig. 2c); other pores seen in the same section, however, ap- pear to be devoid of large cells or connective tissue. Interzooidal communication combined with the ability to secrete protective calcareous diaphragms, was probably one factor in the success of post-Paleozoic cyclostomes. In most Paleozoic bryozoans, the secretion of diaphragms within a zooecium formed a series of closed chambers. Living soft tissues were confined to a zone at the periphery of the zoarium, defined and underlain proximally by the last-formed diaphragm. This skeleton would seem to have provided only a supporting function analogous to that of a coralline calyx with a relatively small protective potential. In post-Paleozoic cyclo- stomes, however, living tissues capable of metabolic activities could be supported throughout the zoarial framework because of the com- munication system of interzooidal pores. This tissue may provide a temporary internal reservoir in time of stress when external condi- tions might be unfavorable to the existence of most feeding zooids. Under more favorable conditions, these underlying tissues might support the proliferation of new feeding zooids allowing the sur- vival of the cyclostome colony. Thus, post-Paleozoic bryozoans, pro- 38 BULLETIN 291 vided with interzooidal communications, have a flexibility in re- acting to environmental changes not possessed by most Paleozoic bryozoans. ZOARIAL BROOD CHAMBERS Occurrence and taxonomic importance of zoarial brood cham- bers have been discussed by Waters (1890), Harmer (1896), Canu (1898, 1899, 1918, 1919a, b), Canu and Bassler (1930), and Borg (1926a, 1933, 1944). Borg (1933, pp. 267-9, pl. 2, figs. 1-3) demon- strated the presence of larvae in typical brood chambers of Recent cerioporid species. In cerioporids, zoarial brood chambers are large skeletal cavi- ties found in exozones (PI. 1, fig. 1f; Pl. 3, figs. 1c, 3; Pl. 12, fig. 2c; PI. 15, figs. 1c, e, f; Pl. 32, fig. 1f; Pl. 34, fig. 2b). General structures are illustrated in Text-fig. 5. The chambers have more-or-less com- plete floors which are compound walls and which seal off most sub- jacent zooecia. The lateral walls are compound walls shared with adjacent zooecia. The chambers are commonly covered by a porous calcareous roof which is similar in appearance to the terminal dia- phragms (PI. 6, fig. 3c; Pl. 33, fig. 1; Pl. 40, figs. 1c, d, e). The nature of the membrane exterior to the brood chamber roof is presently unknown. Borg (1926a, p. 407) stated only that the calcified roof is ““ .. a cryptocyst beneath the original thin gymnocyst”; thus, the pores through the roof are not called pseudopores. If it could be shown that pores are directly sealed by cuticle, then the pores are pseudopores in the sense of Borg. Following abandonment, brood chambers are commonly submerged beneath basal layers which ad- vance from the compound lateral wall and from which new zooecia bud and grow (PI. 3, fig. 1c; Pl. 32, fig. 1f; Pl. 33, fig. 1). Coscinoecia retains a continuous skeletal opening to the surface for some time after it is initially overgrown. In most other cerioporid genera, how- ever, brood chambers are completely submerged. In some genera, zooecia pass through the interior of the cham- ber, appearing like supporting columns. These intrachamber zooecia often have thin-walled septate partitions which extend laterally from them.(P1..32,, fie. 1f Pl. 33.¢fie? 15 0Pls. 51, tig. 1). Whe jpartitions commonly radiate away from the central open area. The intra- chamber zooecia have thin compound walls. These walls were formed 39 CErIoporip CycLostomes (Bryozoa): NYE “sau0}soj9AO PIJodolsad Ul SiaquivYys pooq pauOpuEgE Jo aljoid aysodwod ‘¢ a1NBIz-1xX9 L, is i dd] ac Ohdh anaa¢ VM | 1°a °Y "S°uD' ° 40 BULLETIN 291 by back-to-back deposition from the zooidal membrane and the membrane lining the brood chamber. The septate walls are thinner, and were probably deposited by the brood chamber membrane only (Text-tig. 5, Pl.3Z, fig. LiPl..33; tie. 1). Borg (1933, pp. 269-70) believed that brood chambers were formed by the secondary resorption of the walls of zooecia adjacent to a fertile zooecium. If the chambers had arisen through resorption, then the floor and wall of the brood chamber would have been plas- tered over subjacent zooecial walls unconformably. Thus, micro- structures of the zooecia and the brood chambers should be discon- tinuous and marked in thin-section by a sharp boundary zone. The brood chambers observed in this study, however, are primary struc- tures. The floor and lateral walls of the brood chambers are com- pound walls and structurally continuous with subjacent or adjacent zooecial walls( Pl"6; fig. 3c; Pl AZ, figs. 2d,e; Pl. 15, fig. Ness laaae fig. 1; Pl. 40, fig. 1d). In addition, pores are sometimes seen to con- nect brood chambers to adjacent zooecial chambers. These pores ap- pear in all respects as true interzooidal pores formed as primary structures at the time the wall was deposited. The compound growth of the partitions bounding the brood chamber indicates that the chamber should be considered as a zoarial structure rather than an inflation of a single fertile zooid (gonoe- cium). The zoarial membranes at apertures of numerous zooecia acted in concert to deposit the floor. In Coscinoecia, poorly-preserved microstructure suggests that the floor may have been formed by the progressive ringlike growth of a terminal zooecial closure (PI. 15, fig. lh). In some zooecia, intermediate diaphragms were secreted sub- jacent to the floor (PI. 15, fig. 1h). In other genera, the floor was initially a single continuous closure and deposition continued on both sides. SYSTEMATIC DESCRIPTIONS Order CYCLOSTOMATA Busk, 1852 Suborder CERIOPORINA von Hagenow, 1851 Genus CERIOCAVA d’Orbigny, 1854 Type species: Millepora corymbosa Lamouroux, 1821, p. 87, pl. CERIoPoRID CycLosTomEs (Bryozoa): NYE 41 83, figs. 8, 9 by subsequent designation, Gregory (1896, p. 162). 1821. Pars Millepora Lamouroux, Exposition Méthodique des Genres de |’Ordre des Polypiers, des Zoophytes d’Ellis et Solander, p. 87, pl. 83, figs. 8, 9. 1849. Non Monticulipora d’Orbigny, Rev. et Mag. Zoologie, vol. 1, ser. 2, p. 503. 1854. Ceriocava d’Orbigny, Terrain Crétacé Bryozaires: Paléontologie Fran- caise Description des Animaux Invertébrés, vol. 5, p. 1016. 1896. Ceriocava d’Orbigny, Gregory, Catalogue of Fossil Bryozoa in the Depart- ment of Geology, British Museum (Natural History), The Jurassic Bryozoa, p. 162. 1922. Ceriocava d’Orbigny, Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 61, peo! 1935. Monticulipora d’Orbigny, Bassler, Fossilium Catalogus, I, Pars 67, Bryozoa, pp. 14, 69. 1953. Ceriocava d’Orbigny, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G70. 1953. Non Monticulipora d’Orbigny, Bassler, Treatise on Invertebrate Paleon- tology, Part G, Bryozoa, p. G70. Tentative dtagnosis.—Zoaria_ branching; branches having strongly differentiated coaxial endozone and exozone. Zoarial sur- face smooth to monticular. In endozone, zooecial cross sections markedly small relative to those in exozone. Basal diaphragms widely spaced. In exozone, zooecial walls indistinctly to distinctly laminate. Laminae commonly forming broadly rounded to V-shaped patterns pointing orally. Laminae merging obscurely with structurally in- distinct tissue at zooecial boundary zone. Basal diaphragms more numerous and closely spaced than in endozone. Terminal dia- phragms common. Taxa included.—Only C. corymbosa Lamouroux, the type species, and C. multilamellosa Canu and Bassler were studied. C. multilamellosa was designated the type species of Dendroecita by Cotillon and Walter (1965) and is reassigned here to Haploecia be- cause of the occurrence of peristomial diaphragms, mode of growth, and zooecial wall structure. The internal characters of other species assigned to Ceriocava are unknown to me. Discussion. — D’Orbigny listed 13 species in Certocava with no indication as to which were typical. Gregory (1896, p. 162) desig- nated C. corymbosa as the type species because it is the first recog- nizable species in D’Orbigny’s list, and because it is the oldest and best known species. Bassler (1934, p. 408) discovered that D’Orbigny (1854) had 42 BuLLeTIN 291 designated Monticulipora “frustulosa” (Michelin) (a misspelling for the trivial name pustulosa Michelin, 1846) as the type species of Monticulipora (also Utgaard and Boardman, 1965, p. 112). Bassler (1935, p. 151; 1953, p. G70) listed Ceriopora pustulosa Michelin, 1846 (= Monticulipora “frustulosa” d’Orbigny, 1854) as synonymous with Millepora corymbosa Lamouroux, 1821 (the placement of Ceriopora pustulosa Michelin in synonymy under Millepora corymbosa Lamouroux is not here evaluated because of the lack of information concerning internal characters of Michelin’s species). Unfortunately, Bassler (1935) also listed Certopora pustu- losa as the type species of Ceriocava. This last action is illegal under provisions of the ICZN because it violates priority in view of Gregory’s earlier (1896) designation of Mullepora corymbosa Lamouroux as the type species. Secondly, Bassler’s designation vio- lates provisions concerning availability because Ceriopora pustulosa did not appear in D’Orbigny’s original list of 13 species of Ceriocava, and deliberately so since he had already designated it as the type species of Monticulipora. ‘ Because of his belief that Ceriopora pustulosa was the type species of both Monticulipora and Ceriocava, Bassler (1935, p. 151) cited Ceriocava as a junior synonym of Monticulipora. In 1953 (p. G70) and without explanation, Bassler reversed his earlier position and listed Monticulipora as an objective synonym of Certocava. At the petition of Bassler and Duncan, the ICZN (1955, Opinion 443) designated Monticulipora mammulata dOrbigny, 1850, as the type species of Monticulipora d’Orbigny, 1849. Thus at present, there is no justification for placing Monticulipora in synonymy with Ceriocava or vice-versa, with or without regard to the conspecificity of Ceriopora pustulosa Michelin and Ceriocava corymbosa Lamouroux. D’Orbigny established the genus Ceriocava in order to dis- tinguish “... tous les Ceriopora des auteurs ayant une seule couche de cellules et des ouvertures simples, représentant dans leur ensem- ble, une colonie rameuse” (1854, p. 1015). Gregory (1896, p. 163) emphasized the “ . . . thick, irregularly branching habit . . .” and added information on some internal characters: “The axis of the zoarium consists of fine zooecia densely packed. The outer zone consists of zooecia which are usually reflexed and of much greater CERIOPORID CycLosToMEs (Bryozoa): NYE 43 diameter.” Gregory illustrated a specimen of C. corymbosa, the type species, with numerous diaphragms in the endozone (text-fig. 13, p. 164). Canu and Bassler (1922, pp. 90-2, text-fig. 20, p. 91) also described and illustrated thin sections of this species. The illustrations clearly show numerous diaphragms in the endozone, and an increase of zooecial diameters in the exozone relative to the endozone. One character not mentioned or illustrated is the occur- rence of basal diaphragms in the exozone. Ceriocava differs from Coscinoecia in having basal diaphragms in the exozone, and in having a V-shaped laminar structure and light-colored tissue in the outer exozone which often narrowly out- lines the zooecial boundary zone. These genera differ in other charac- ters such as details of brood chambers, architecture, and occurrence of mural spines; but these characters are tentatively considered diagnostic at the specific level. Ceriocava differs from Reptonodicava in having branched zoaria with well-differentiated coaxial exozone-endozones, and in wall structure. Ceriocava corymbosa (Lamouroux), 1821 Pl. 1,, figs. Ta-h; Ply 2, figs. la-h; Pl. 3, figs. la-c, 2, 3; Pl. 4, figs. la-e; Pl. 5, figs. la-c, 2, 3; Pl. 6, figs. 1, 2, 3a-c 1821. Millepora corymbosa Lamouroux, Exposition Méthodique des Genres de l’Ordre des Polypiers, des Zoophytes d’Ellis et Solander, p. 87, pl. 83, figs. 8, 9. 1854. Ceriocava corymbosa (Lamouroux), d’Orbigny, Terrain Crétacé Bryo- zoaires: Paléontologic Francaise Description des Animaux Invertébrés, vol. 5, p. 1016. 1896. Ceriocava corymbosa (Lamouroux), Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural Histry), the Jurassic Bryozoa, pp. 163, 164, text-fig. 13, p. 164. 1922. Ceriocava corymbosa (Lamouroux), Canu, and Bassler, U.S. Nat. Museum, Proc., vol. 61, p. 90, pl. 14, figs. 5, 6, 8; p. 91, fig. 20. 1965. Ceriocava corymbosa (Lamouroux), Cotillon, and Walter, Soc. Géol. France Bull., vol. 7, ser. 7, p. 935. Type. — Lamouroux’ specimens of C. corymbosa, originally stored in the Museum at the Université de Caen, were probably destroyed during the invasion of Normandy in World War II (fide Prof. L. Dangeard, Université de Caen). Lamouroux did not desig- nate a holotype, and no specimen is known to have been designated as the lectotype. 44 BULLETIN 291 Type locality and horizon. — “Terrain a polypiers environ de Caen” (Jurassic, Bathonien, Calvados, France). Material studied. — Thin sections and acetate peels were made from the following topotypes: MNHN IP2-1, IP2-2, Bathonien, Ranville (Calvados), France; USNM 32164-1, -2, -3, Bathonien, St. Aubin (Calvados), France; USNM 32181-1, -2, -3; USNM 68941-1, -2, Bathonien, Ranville (Calvados), France. USNM 68941-1 was figured by Canu and Bassler, 1922, p. 14, fig. 8 and USNM 68941-2, pl. 14, fig. 6. Duplicate acetate peels are preserved in the National Museum of Natural History, Washington, D.C., and the author’s collection. Description. — Growth habit. — Branches are robust, and have subcircular to elliptical cross sections. Monticules are ridgelike to pustulose. Intra- zoarial overgrowths occur, but are generally local in extent. The exozone and endozone intergrade in a broad zone of zooecial bending. Endozonal zooecia grow approximately parallel to the major branch axis for relatively long distances (PI. 4, fig. le). Orally from the zooecial bend, zooecia are commonly rectilinear in growth and intersect the zoarial surface at approximately 90°. Branches are massively intergrown in axillary zones distal to the bifurcation of branches (PI. 2, fig. 1h; Pl. 4, fig. 1b). In the axillary zone, zooecia in each new branch are recurved distally as the distance separating the growth surfaces of each branch approxi- mates 1 mm. Distally from this axillary zone of bending, the zooecial walls intersect obliquely. At the intersection the walls anastomose, forming a continuous compound wall against which zooecial chambers are pinched out (PI. 4, fig. 1b). Zones of irregular zooecia were occasionally observed. In one mode, zooecia are small in diameter, thin-walled, and irregular in growth direction (PI. 3, figs. 1b, 2). In a second mode (PI. 3, fig. la), the zooecia bud from a diaphragm-like structure, sealing off subjacent zooecial cavities. Superjacent zooecial walls are struc- turally continuous with the diaphragm-like structures. Endozone — Zooecial walls show cyclic repetition of structure (Pl. 1, fig. 1d; PI. 2, fig. 1h). In each cycle (PI. 4, fig. le), individual zooecial walls are thin and straight to slightly undulatory for a rela- CerioporID CycLosToMEs (Bryozoa): NYE 45 tively long distance. Distally, marking the boundary of the cycle, zooecial walls are thickened annularly and form a zone parallel to the growing tip of the branch. Zooecial cross sections are small and elliptical to subelliptical (PI. 2, fig. 1f). Interzooidal pores were rarely observed. Zooecial walls are dark in color and indistinctly granular with a thin zooecial lining. Basal diaphragms (PI. 5, fig. 2) are thin (less than .0016 mm), dark in color and commonly convex aborally, and they flex orally to merge with the zooecial lining. Exozone — Zooecial walls are generally symmetrical (less com- monly slightly asymmetrical) in thickness across the zooecial boundary zone (PI. 1, fig. 1g; PI. 5, fig. 1c). Zooecial walls generally have distinctly moniliform profiles due, in part, to the occurrence of numerous, large and widely flared interzooidal pores (PI. 5, fig. le; Pl. 6, fig. 3b). Moniliform profiles are commonly oblate to elliptical near the zooecial bend (PI. 3, fig. 1a), becoming alate to sagitate orally (Pl. 3, fig. 1c; Pl. 4, figs. 1c, d). Zooecial chambers are com- monly subcircular in cross section. Mural spines are abundant (PI. 5, fig. 3; Pl. 6, fig. 2) to nearly absent (PI. 1, fig. 1h). Diaphragms — Terminal diaphragms are numerous. The dia- phragms are thick (Table 4, TrID-Th) and are commonly seen to occur slightly aboral to the skeletal aperture (PI. 1, figs. 1d, f; PI. 2, figs. lle, f; Pl. 5, fig: la; Pl. 6, fig. 3a). Ihe oral surfaces of the diaphragms are generally planar, but the aboral surfaces are often uneven (PI. 5, fig. la). The diaphragms sometimes adjoin the zooecial wall with slight flexure, or have relatively thick aborally flexed abutments (PI. 5, fig. 1a; Pl. 6, fig. 3a). Diaphragms occasion- ally show slight flexure towards the aperture (PI. 5, fig. la; proxi- mal portion of top diaphragm). Basal diaphragms are numerous and closely spaced (see Table 4 BasiOsint; Pil 1, figs Lis Plt 2) tie. Ve; Pl? 3) fre Ta) Phe dia- phragms are thin (less than .0016 mm) and slightly convex aboral- ly, and they merge obscurely with the zooecial lining (BES; ties fe; EP G.shigss Jassb))). Brood chambers — Roughly oblate brood chambers were ob- served in the exozone (PI. 1, fig. 1f; Pl. 2, fig. 1g; Pl. 3, fig. lic): The chambers are large; measurements of the longest dimension parallel to the zoarial surface (BrCh-Wth) and of the dimension 46 BULLETIN 291 normal to the zoarial surface (BrCh-Dth) are given in Table 4. The floor is complete but is pierced by occasional interzooidal pores (PI. 6, fig. 3c). The porous roof is thick (Table 4, BrChRf-Th). Aban- doned brood chambers are submerged beneath a thin basal layer. Zooecia budding from the basal layer are either thick-walled with moniliform profiles and relatively straight in growth, or they are thin-walled, parallel-sided and irregular in growth (PI. 3, fig. Ic). Discussion. — Unfortunately, the external characters of Cerio- cava corymbosa as figured by Lamouroux are not diagnostic. Later authors have referred to C. corymbosa, but none had redescribed or refigured the type specimens prior to their presumed destruction (see “Type” above). Under such circumstances, the assignment of specimens to Lamouroux’ species and continued usage of the name is open to some question. The species concept, however, is reasonably consistent from author to author as seen in the descriptions and illustrations of Gregory (1896, pp. 164-5, text-fig. 13, p. 164), Canu and Bassler (1922, pp. 90-92, text-fig. 20, p. 91, pl. 14, figs. 1-8; 1929, p. 115, pl. 1, figs. 4, 5) and Cotillon and Walter (1965, p. 935). Furthermore, specimens examined by Gregory and Canu and Bassler were collected from the type locality, and the authors cited above considered internal characters in their diagnoses. For these reasons, and for the purpose of stability, the continuation of C. corymbosa (Lamouroux) seems justified. Remarks concerning mode of growth. — Zoaria of C. corymbosa exhibit the typical coaxial exozone and endozone mode of growth of most large ramose Stenolaemata. Also, the periodic annular thickening of the zooecial walls in the endozone is essentially similar to cyclic growth phenomena demonstrated in other stenolaemates, such as in three living cerioporid species (Borg, 1933), two Devonian trepostome species (Boardman, 1960) and in the Upper Paleozoic trepostome, Tabulipora Young (Gautier, 1970). Intrazoarial over- growths were observed only in localized areas, and do not appear to play a major role in zoarial increase. One specimen of C. corymbosa was found which completely encrusts a branch fragment of Hap- loecia straminea (Phillips). The formation of an axillary zone of intergrown zooecia pre- sumably functions to strengthen the colony. The recurvature and anastomosing growth of zooecia is not unique to C. corymbosa but 47 Cerioporip CycLosTomEs (Bryozoa): NYE S19JIUIT|[TW UT, $ I $ 4 st 200° $T0° T10° 610° T1QUN-1dPZ I 9 OSZ Ose 89 9¢ se 0 Ove ugspoZ/UD-dsoZ i 9 OSz OSZ 0 0's US§097/FD=dPZ I 9 Ose OSc Le 20° 80° £0° +1" YL-IMPZPO UQX*INN-9S8)-4D9Z I 9 0SZ 0S2 at c ct OT Ve UQXIN-US8D-409Z I Y) 0Sz Osc at £0° £c° It Og UQXWN-YS89-4D9Z T 9 Osc OSc at +0" 8c" +r 8e° UGS INsTSS OP JUOZOXY - [B199007 L L LE Le HL 6¢ SIT UGE SSsOs a [el1e07 2poD ‘vadg IZN ZN N ‘A‘O #S 4X «XO EARLE AAG) (XQNOUYNOWNVT) FSOIWANOD FAFIOINAD AO SLNAWANNASVAW JO AUVWINDS IVOLLSILLV.LS + ATEVL BULLETIN 291 48 SI9}OUMN|[TW UT UQXxW-4Ss)-7d GIA.L QL-dIAL auo0zoxg - wseiydeiq GL-FTIM9Z W$899Z/UD-1dPZ YL-IM9PZPO UQXWN-4S8)0-4YD9Z UQXW-4Ss)-YO2Z UCXIAN-USSD-Y99Z UQXI-USsD-YOIZ auozopuy - [v199007 S I 8I £2 400° LT0° Z10° $z0° ) £ 12 9b Z0° $0’ Z0° 4 £ c Os Os 0 £00° c £ SL SZ 0 07 c £ SL SL st 10° ZO" T0° +0" c £ SZ SZ 8T (a eT OT V7 c £ SZ SZ 14 Z0° L0° ZO" cr c £ $Z SZ 1d Z0° 60° £0° A apoD ‘sadg IZN %ZN N “AO #S aX # LO 1a}OBIEYO (XNOWNOWVT) FSOIWANOD FAFIOINAD AO SLNAWAANSVAW AO AUVWWAS TVOILSILV.LS 49 Certoporip CycLostomes (Bryozoa): NYE ‘C-b91ZE WNSD ‘2-@dI NHNW ‘T-@dI NHNW ‘sadAjodoy, ° ‘I-ZdI NHNV :sedAjodo 7, ° (Sc) Z-181Z€ WNSA ‘(S2) @-%dI NHNW ‘(S2) I-@dI NHNW :82d430d0 7, ($2) Z-1+689 WNSA ‘(SZ) 2-+91ZE WNSN :sedAjodoy, ° "(SZ) Z-1b689 WNSN ‘($Z) 2-491ZE WNSA ‘(SZ) @-ZdI NHNW :sed4jodoy, ° "(S$Z) Z-P91ZE WNS?N ‘(SZ) 2-1+689 WNSNO ‘(SZ) @-18TZE INNSO ‘(SZ) I-I8IZe WNSQA ‘(001) 2-ZdI NHNW ‘(0S) I-ZdI NHNIW :2d4}0doy, ‘1 ad0O0 NAWIOddS OL AAW Non tw s SI9JIUIT[ [IW UT, $ I I Or 50° st It 9S" WLFAO $ I 6 te 60° Ue +0 or YL-1d4O1d § I zZ ¢° ¢ Wa-Yo!d $ I Z ; Ms WAA-YOE Iaquiryd pooig v £ ST SL Ad 80° tc 80° 0S" quyJ-d[sa (‘uod) suozoxy - wSeiydeig apo neds = ZN OOZNONN AD «8 «X «WO sayoereYyO (XNOYNOWVI) FSOFWANOD FAPIOINAD AO SLNAWAYNSVAW AO AUVWWAS TVOLLSILVILS 50 BULLETIN 291 TABLE 5 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN CERIOCAVA CORYMBOSA (LAMOUROUX)* Cire. Ellip. Ov. Pyr. Polyg. Triang. Irreg. Exozone Regular 1 Sub 142 38 17 Irregular 24 22 + Endozone Regular 14 3 3 Sub 2 25 12 2 1 Irregular 1 10 2 *Estimates of exozone cross-sections were made from MNHN IP2-1 (50 zo- oecia), IP2-2 (100 zooecia), USNM 32181-1 (3 zooecia), USNM 32181-2 (25 zooecia), USNM 68941-2 (25 zooecia) and USNM 32164-2 (25 zooecia). Endozonal zooecial shapes were estimated from MNHN IP2-2 (25 zooecia), USNM 32164-2 (25 zooecia) and USNM 68941-2 (25 zooecia). was observed at the oblique intersection of distally growing branches of Haploecia multilamellosa (Canu and Bassler) and Parletosoecta jacksonica Canu and Bassler. R. S. Boardman (pers. comm. 1971) stated that zooecial recurvature is typical of trepostomes in the same situation. The recurvature of zooecia suggests that the initial stimulus for change in growth habits is related to the proximity of zooecia grow- ing towards each other from each new branch. Possibly the stimulus is provided by direct contact of soft tissues. Recurvature, however, begins when zooecia are separated by as much as 1 mm; thus, if direct contact provides the stimulus, contact would be limited to the tips of tentacles. The mode of anastomosis is similar to that seen in Parleiosoecia jacksonica in that zooecia growing towards each other tend to be- come recurved, and to approach the surface of contact obliquely. In both species, zooecia form compound walls of anastomosis with opposing zooecia, and individual zooecia are pinched out as growth continues. Anastomosis of zooecia, however, commonly occurs at the juncture of two distally growing branches in P. jacksonica rather than following the bifurcation of a single branch into two branches. Also in P. jacksonica, the zooecial walls of intersecting zooecia be- CERIOPORID CycLosTOMES (Bryozoa): NYE 51 751A 75 B ~> > oO oO C= S50 250 o rom g 25 25 mm 4a 22 a6 1.0 2.0 ZcCh-CsSn-NMxDn ZcCh-CsSn-MxDn ZcCh-CsSn-NMxDn 751C > Fé 100 c o 3 90 xs 75 rox © 2 — = 50 25 x p= | = 25 =) oO mm A 32. 1 2.3 4 5° 6 CdZcWI-Th ZdPr-Cn/ZcCsSn Text-figure 6 A-D. Histograms and cumulative curve from three topotypes of Ceriocava corymbosa (Lamouroux). A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness. D. Count of interzooidal pores per zooecial cross section. EM BULLETIN 291 come thin-walled and parallel-sided prior to merger. In C. corym- bosa the zooecial walls remain thick-walled and have moniliform profiles up to the merger of intersecting zooecial walls. In addition, in C. corymbosa the new wall formed at the zone of intersection is moniliform in profile and thick-walled (PI. 4, fig. 1b). Irregularities in budding habit and zooecial appearance occur locally in the exozone in C. corymbosa. In one mode, new zooecia are budded from thick diaphragm-like structures which seal off sub- jacent zooecia (PI. 3, fig. la). In a second mode, normally appearing zooecial walls terminate and are followed by irregularly oriented, thin-walled zooecia. Although the zooecial chambers appear to be continuous from normal to irregular zooecia, the zooecial walls are discontinuous and the walls of irregular zooecia are continuous with adjacent zooecial walls (Pl. 3, fig. 2). These irregularities are inter- preted as a specialized zoarial response to disease or trauma by which zoarial growth may proceed. If so, this differs from other species in which zoarial repair is affected by the extension of a basal layer over dead portions of the zoarium, e.g., Reptonodicava (PI. 43, figs. 3a, b; Boardman, 1960, p. 39). Genus CERIOPORA Goldfuss, 1826 Type species: Ceriopora micropora Goldfuss, 1826, p. 33, pl. 10, figs. 4a, d, by subsequent designation, Gregory (1896, p. 195). 1826. Pars Ceriopora Goldfuss, Petrefacta Germaniae, vol. 1, p. 32. 1830. Pars Ceriopora Goldfuss, Blainville, Zoophytes: Dictionnaire de Science Naturelles, vol. 60, p. 378. 1834. Pars Ceriopora Goldfuss, Blainville, Manuel d’Actinologie ou de Zoo- phytologie, p. 413. 1851. Pars Ceriopora Goldfuss, von Hagenow, Die Bryozoen der Maastrichter Kreidebildung, p. 52. 1851. Pars Heteropora Blainville, von Hagenow, Die Bryozoen der Maastrichter Kreidebildung, p. 52. 1896. Pars Ceriopora Goldfuss, Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Juras- sic Bryozoa, p. 195. 1909. Pars Ceriopora Goldfuss, Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Cre- taceous Bryozoa, vol. 2, p. 156. 1953. Ceriopora Goldfuss, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G57. 1953. Non Reptonodicava d’Orbigny, Bassler, Treatise on Invertebrate Paleon- tology: Part G, Bryozoa, p. G57. Tentative diagnosis. — Zoaria massive with radial growth habit modified by development of major growth axis. Zoarial increase CrERIopoRID CycLosTomeEs ( Bryozoa): NYE 53 in part of repetitive, but irregularly occurring, intrazoarial over- growth. Endozone commonly restricted to portions of zooecia just adjacent to basal wall of intrazoardial overgrowths. In exozone, zooecial walls exhibiting variable repetition of thin-walled and thick- walled phases. In thin-walled phase of exozone, zooecial wall granular to in- distinctly laminate. In thick-walled phase, zooecial walls distinctly laminate, laminae arching orally convex, continuous across zooecial boundary zone. Light-colored granular tissue commonly forms small, rounded masses in outer cortex. Intermediate diaphragms occurring in zooecia subjacent to in- trazoarial overgrowths. Taxa included. — Based on observations of internal characters seen in the primary types and identified specimens from the type localities, the following species originally included in Certopora by Goldfuss are considered to be correctly assigned to other genera: Ditaxia anomalopora (Goldfuss), Heteropora cryptopora (Goldfuss ) and Zonopora spiralis (Goldfuss). Based on observations of the internal characters seen in identified specimens from the type locality, C. globosa Michelin is considered to be correctly assigned to Reptonodicava. Based on the observation of internal characters seen in identified specimens, C. dichotoma Goldfuss is tentatively referred to as Grammascoecia. The internal characters of other species assigned by others to Ceriopora are unknown to me. Discussion. — Goldfuss (1826, p. 32) erected the genus Certo- pora and assigned 28 new species to it. He did not designate or in- dicate a type species. Gregory (1896, p. 195) listed the type species as C. micropora Goldfuss without comment, but later (Gregory, 1909, pp. 156-7) he indicated that his designation of lectotype fol- lowed, in part, the previous restrictions of Blainville (1830) and D’Orbigny (1854). The concept of Ceriopora was based largely on two external characters (see Gregory, 1909, pp. 156-158 for an excellent review of development of the generic concept in the last century): 1) Aperture size — Blainville (1830, p. 378) separated Heteropora from Ceriopora. He stated that two kinds of zooecia could be recognized in Heteropora. Each polymorph was characterized by 54 2) BULLETIN 291 the size of its aperture. Thus, Blainville implied that only one kind of zooecium could be recognized in Ceriopora. This generic distinction was followed by D’Orbigny (1854, p. 1029), Gregory (1896, p. 195; 1909, p. 156), Canu and Bassler (1920, p. 678), and Bassler (1953, p. G67). The concept of one kind of zooecium based on the size of the aperture was referred to as “mono- morphic”, or “lacking mesopores” by authors after D’Orbigny. The continued use of this character as generically diagnostic does not appear to be justified, however, because the frequency dis- tributions of cross-sectional dimensions of zooecial chambers made from the type specimens of both Ceriopora micropora Gold- fuss and Heteropora cryptopora (Goldfuss), their respective type species, are unimodal, and approach a normal distribution (Text- figs. 7A and 14A). Intrazoarial overgrowth — species with overgrowths have been referred to as “multilamellar” (Canu and Bassler, and Gregory), or “plusiers couches superposées” (d’Orbigny). Overgrowths can often, but not invariably, be recognized on the external surfaces of zoaria, especially when the surface is slightly worn. De Blain- ville (1830) and D’Orbigny (1854) believed that zoaria of Ceriopora were composed of several overgrowths. D’Orbigny’s earlier position was followed by Canu and Bassler (1926, p. 19) and Gregory (1909, p. 157). Gregory noted that “a certain amount of marginal lamellation must be expected in massive Bryozoa”. Ceriopora, as based on observations of the type speci- mens, does have intrazoarial overgrowths and would thus be multilamellar in the sense of Blainville and D’Orbigny (1854). The mode and amount of intrazoarial overgrowth is apparently a useful taxonomic character. The mode of growth shown in the type species of Ceriopora is intermediate in appearance between Reptono- dicava and Heteropora. The type species of all three genera show a reduced endozone, apparently restricted to the proximal-most por- tion of encrusting zooecia, budding from a basal layer. All three are erect, massive, and globular to subramose in appearance. In Hetero- pora, intrazoarial overgrowth plays an important role in zoarial in- crease. The zoarium is composed of nested intrazoarial overgrowths connected by a few zooecia with continuous zooecial chambers, but Certoporip CycLosTomeEs (Bryozoa): NYE 55 the zooecial walls from subjacent to superjacent growth commonly show some evidence of discontinuity of growth, such as great reduc- tion in size and separation by a dark line. In the type species of Reptonodicava overgrowths are rare, covering only a few zooecia at most, and zooecia commonly grew continuously in all directions for very long distances. In the type specimen of Ceriopora intrazoarial overgrowth plays an intermediate role relative to Heteropora and Reptonodi- cava. Gregory (1896, p. 95) included the statement, “Diaphragms horizontal, numerous”, in his diagnosis of Ceriopora. This inclusion was based on his study of thin sections of Jurassic specimens which he assigned to Ceriopora globosa Michelin. The internal characters of C. globosa Michelin are sufficiently different from those of C. mic- ropora to separate both in different genera. Bassler (1935, p. 186) designated C. globosa Michelin as the type species of Reptonodicava, but indicated that Reptonodicava was a synonym of Ceriopora. R. globosa does have numerous, closely spaced basal diaphragms, and the reference to numerous diaphragms in the definition of Ceriopora was reiterated by Canu and Bassler (1920, p. 678) and Bassler (1953, p. G67). Diaphragms occur in the lectotype of C. micropora, but they are intermediate diaphragms and no more than one was observed to occur within a single zooecium, unlike the numerous basal diaphragms typically seen in R. globosa. Remarks on wall structure.— Light-colored, optically nearly structureless tissue in the zooecial wall is interpreted as being orig- inally granular. The cortex of the zooecial walls in the endozone and thin-walled exozone portions are composed almost completely of granular tissue. Lamination becomes increasingly distinct in the thick-walled portions, and light-colored tissue is restricted to dis- continuous bodies in the outer cortex alternating with, and sur- rounded by, laminated tissue. Masses of light-colored tissue bounded conformably by laminate tissue probably represent originally granu- lar tissue (PI. 9, fig. 1b — lower, thin-walled portion of zooecial wall in center — Ic). Masses of light-colored tissue which cut across laminae are inferred to be recrystallized from originally (at least in part) laminate tissue (Pl. 9, fig. 1b — upper, thick-walled por- tion of zooecial wall in center). 56 BULLETIN 291 The wall structure is similar to that seen in Coscinoecia radiata Canu and Lecointre (PI. 14, figs. le, f; Pl. 15, fig. 1g). Ceriopora micropora Goldfuss, 1826 Pl. 7, figs. la-f; Pl. 8, figs. la-d; Pl. 9, figs. la-d 1826. Pars Ceriopora micropora Goldfuss, Petrefacta Germaniae, vol. 1, p. 33, pl. 10, figs. 4a, d; not figs. 4b, c. 1830. Ceriopora micropora Goldfuss, Blainville, Zoophytes: Dictionnaire de Science Naturelles, vol. 60, p. 378. 1834. Ceriopora micropora Goldfuss, Blainville, Manuel d’Actinologie ou de Zoophytologie, p. 413. 1851. Pars Ceriopora micropora Goldfuss, von Hagenow, Die Bryozoen der Maastrichter Kreidebildung, p. 52, pl. 5, fig. 13. 1909. Pars Ceriopora micropora Goldfuss, Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Cretaceous Bryozoa, vol. 2, p. 158-161. 1953. Pars Pennipora beyrichi Hamm, 1881, Voigt, Geol. Staatsinst. Hamburg, Mitt., Bd. 22, pp. 58, 61, 62, pl. 2, fig. 4. Type. — UB 119 is designated as the lectotype. This specimen was figured by Goldfuss (1826, pl. 10, figs. 4a, d), von Hagenow (1851, pl. 5, fig. 13 as Heteropora crassa von Hagenow, 1851), Voigt (1953, pl. 2, fig. 13 as Pennipora beyrichi Hamm, 1881), and here (PI. 7, figs. la-f; Pl. 8, figs. la-d and PI. 9, figs. la-d). Type locality and horizon. —“St. Petersberge bei Maastricht [Limbourg, Netherlands], aus dem Mergel bei Essen an der Ruhr [Federal Republic of Germany], und aus der Conchilienbreccie in der obern Schicht der Kreide von Cleom bei Nantu”. Locality data were not listed with the figures, and Von Hagenow (1851) ap- parently did not find any locality data with the specimens. Com- monly, Ceriopora micropora is listed as a Maastrichtian fossil from Maastricht, Netherlands. Many specimens from the area of Maastricht, externally similar to the lectotype, were sectioned. On the basis of internal characters, these specimens were assigned to Heteropora cryptopora or to two other species referable to other genera perhaps unnamed at present. Thus, reference of C. micropora to Maastrichtian at Maastricht, Netherlands, should be considered questionable. Material studied. —The lectotype UB 119 was borrowed from Geol.-Paleont. Inst. Bonn University, West Germany. Three thin sections and three acetate peel replicas on a single acetate slide were prepared. Much of the specimen remained after sectioning. Dupli- CertoporIp CycLosToMEs (Bryozoa): NYE 57 cate acetate peels are preserved in the National Museum of Natural History and the author’s collection. Description of the lectotype. — This description is based solely on the lectotype, the only specimen of C. micropora known to be available for study. This description includes assessment of non- genetic variation within this colony. No assessment of genetic or other interzoarial variation within C’. micropora is implied in this description. Endozone — The zooecial walls are undulatory to straight and thin (about .007 and .012 mm). The walls have a granular cortex and a thin, dark zooecial lining (PI. 7, fig. 1d; Pl. 8, figs. 1c, d). Exozone — In the thin-walled exozone phase, zooecial walls are commonly thickened symmetrically across the boundary zone, and are submoniliform in profile. Monili are generally clavate to fusi- form. Interzooidal pores are rare. Zooecial walls are homogeneous to slightly granular with a thin zooecial lining. In the thick-walled exozone phases, zooecial walls are thickened symmetrically to asymmetrically across the boundary zone, and have moniliform cross sections. Successive monili are commonly un- equal in length and thickness (PI. 9, figs. 1b, c), and sometimes show abrupt changes in growth direction among the monili (PI. 9, fig. 1b). Zooecial chambers are commonly subelliptical in cross section, but show great variability (Table 7). Interzooidal pores are numerous (Text-fig. 7D). Diaphragms — Intermediate diaphragms are occasionally ob- served, up to one per zooecium. The diaphragms commonly occur about .1-.2 mm aboral to zooecial apertures, and subjacent to intra- zoarial overgrowths (PI. 9, figs. la, b). The oral surfaces of the dia- phragms are generally planar; the aboral surfaces are planar to strongly convex orally, and sometimes have an aborally flexed abut- ment which tapers into a short zooecial lining (Pl. 9, figs. 1a, b). Diaphragms are highly variable in thickness (Table 6, IntD-Th). 58 BULLETIN 291 751A Oe B > ~ oO oO (= (Ss o 530 25 Ss oO ® ® nN uo Nh oi mm 4d 2. a J 1.0 20 ZcCh-CsSn-NMxDn ZcCh-CsSn-Mx Dn ZcCh-CsSn-NMxDn 751C > re 100 = © 5 90 75 rom ® : a = 50 25 a >} ¢ 25 =} oO mm | Pa 1273’ 456 CdZcWI-Th ZdPr-Cn/ZcCsSn Text-figure 7 A-D. Histograms and cumulative curve from the lectotype of Ceriopora micropora Goldfuss. A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness. D. Count of interzooidal pores per zooecial cross section. 59 CERIOPORID CycLosTOMEs (Bryozoa): NYE ad0O NAWIOddS OL AIM 61 AN 244303997 “T SIDJIWI|[IW Ul» T T a S co O10 910° 800° 0£0° YL-dta] auo0zoxy-wdseiydeig T T IT £T th +00° 600° £00° +10 IGUN-1dPZ T i OOT OOT 0 M4 WS899Z /UD=1dPZ T T OOT Oot ch 810° c+0" 800° Lor YL-LMA9ZPO UQXINN-9S8)-409Z T I oot oot Al Z0 eT OT oc UGXIN-USsD-YO9Z T T Oot oot £7 20° [fs $0° os UGXWN-US80-YOIZ T T OOT oor fc £0° +r $0° to BCXIN=US§O S402 au0ZOxy - [8199007 T T T OT DWM-1Z T I I 02 tH-IZ [Bl1e0Z apop 99dg_ — IZN -OZN N AYO #S 4X «WO CHER EAL LO) SSQOAATOO FAOdOUIDIW FUOdOINAI 9 ATAVL JO SLNANAYNASVANW AO AUVWWOAS TVOLLSILVLS 60 BULLETIN 291 TABLE 7 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN CERIOPORA MICROPORA GOLDFUSS Cires) Bilip: Ov. Pyr. Polyg. Triang. Irreg. Exozone Regular 17 7 Sub 2 38 8 3 Irregular 3 10 2 Endozone . Regular 4 4 Sub 1 10 4 1 Irregular . 1 Discussion. — Designation of UB 119 as the lectotype agrees with restrictions first made by Blainville (1830, p. 378) and, in part, with restrictions in reference to figure 4d, but not figure 4a made by Von Hagenow (1851, p. 52) and followed by Gregory (1909, pp. 159-60) and Voigt (1953, pp. 58-62). Von Hagenow (1851, pp. 48, 51) stated that he had studied three specimens which he believed were the original specimens be- fore Goldfuss when he described Ceriopora micropora. The three specimens were attached to a small tablet (“Tafelchen”) found in the Bonn Museum. Two were syntypic, as they were clearly identi- fiable with the specimens figured by Goldfuss; the first as plate 10, figure 4a — UB 119, and the second as plate 10, figures 4b and c. Von Hagenow was of the opinion that neither of these two specimens was consistent with the concept of C. micropora. He was able to demonstrate that the specimen figured by Goldfuss as plate 10, figures 4b and c was a sponge, and he assigned UB 119 to his new species Heteropora crassa. Von Hagenow believed that the third specimen on the small tablet and Goldfuss’ plate 10, figure 4d were consistent with the concept of C. micropora and referred to both the specimen and the figure as the name-bearer for C. micropora. Furthermore, Von Hagenow believed that Goldfuss had taken the magnified view (pl. 10, fig. 4d) from the third specimen; but as Gregory (1909, p. 160) noted: “Von Hagenow admitted an element of doubt in reference to fig. 4d.” CERIOPORID CycLosToMEs (Bryozoa): NYE 61 Gregory (1909, pp. 159-60) and Voigt (1953, pp. 61-62) ac- cepted Von Hagenow’s restrictions and designation. Voigt (1953, pp. 58-64), however, believed that UB 119 is not conspecific with H. crassa Von Hagenow but is assignable to Pennipora beyrichi Hamm, 1881. These contentions are not here evaluated because of the present lack of knowledge concerning the internal characters of the respective types. Consideration of characters revealed in thin- sections of specimens hitherto considered to be assignable to named species of such genera as Ceriocava, Ceriopora, Heteropora, Repto- multicava, and Reptonodicava, suggests that external characters are generally not diagnostic at the generic and family, and often not even at the species level. For reasons listed below, Von Hagenow’s contention that figure 4d of Goldfuss was taken from the specimen figured by Von Hage- now (1851, pl. 5, fig. 4) is rejected. 1) Goldfuss clearly indicated that plate 10, figures 4a and 4d were made from the same specimen by connecting them with a dashed line. Goldfuss and other authors commonly used the convention of connecting different views of the same specimen by a dashed line. Von Hagenow demonstrated that figures 4b and 4 c of Gold- fuss’ plate 10 were different views of the same specimen; both views were connected by a dashed line. Figures 4a and 4d should be considered as different views of UB 119 unless strong evidence to the contrary were to be offered. The lectotype is close enough in appearance for acceptable com- parison with the magnified and somewhat idealized view of the surface as figured by Goldfuss (pl. 10, fig. 4d). 3) Von Hagenow was not positive of equating the Goldfuss figure (pl. 10, fig. 4d) with the third specimen on the card (Von Hage-~ now, pl. 4, fig. 5), and Von Hagenow emphasized that only the figure by Goldfuss and the third specimen fitted the concept. As outlined above, Von Hagenow is believed to have made a mistake in equating the specimen he figured as plate 5, figure 4, with plate 10, figure 4d of Goldfuss, and thus he designated two specimens to be the name bearer. Von Hagenow’s elimination of the 2 4 sponge specimen (Goldfuss, pl. 10, figs. 4c, d) from consideration is here followed, and the lectotype is designated as the remaining figured specimen, UB 119 figured by Goldfuss (pl. 10, figs. 4a and 4d). 62 BULLETIN 291 Remarks on enigmatic structwres.—A single, unusual dia- phragm-like structure was observed in the transverse section and is illustrated in Pl. 9, figure 1d. This structure, known only from a single profile view, apparently forms a hollow, tubelike structure within the zooecial chamber which flares, and is attached by its oral and aboral extremities to the zooecial wall. The structure is con- sidered to be primary because of its structural continuity with the zooecial wall. Cross sections of two large chambers were observed in tangential section (one is figured in PI. 8, fig. la). The cross-sectional dimen- sions are more than twice as long as the largest zooecia observed (Table 6), and are interpreted as interzooecial structures. The walls are unbroken and are interpreted as primary in origin. The only large primary chambers in other cerioporids are thought to be brood chambers. In lieu of better evidence, these inter-zooecial spaces in C. micropora are interpreted as brood chambers. Genus CORYMBOPORA Michelin, 1846 Type species: Corymbopora menardi Michelin, 1846, p. 213, pl. 53, fig. 10 by monotypy. 1846. Corymbopora Michelin, Iconographie Zoophytologique, description par localités et Terrains des Polypiers Fossiles de France et pays environ- nants, p. 213. 1850. Pars Fasciculipora d’Orbigny (1846), D’Orbigny, Prodrome de Paléon- tologie stratigraphique universelle, vol. 2, p. 177. 1854. Corymbosa d’Orbigny, Terrain Crétacé Bryozoaires: Paléontologie Fran- caise Description des Animaux Invertébrés, vol. 5, p. 691. 1890. Pars Fasciculipora d’Orbigny (1846), Pergens, Soc. Belge Géol. Paleont. Hydro. Bull., vol. 3, p. 377. 1909. Corymbopora Michelin, Gregory, Catalogue of Fossi] Bryozoa in the Department of Geology, British Museum (Natural History), The Cre- taceous Bryozoa, vol. 2, p. 45. 1916. Non Corymbopora Lang, Ann. Mag. Nat. Hist., ser. 8, vol. 18, p. 382. 1919. Corymbopora Michelin, Canu, Soc. Géol. France, Bull., vol. 17, ser. 4, p. 348. 1953. Corymbopora Michelin, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G70. Tentative diagnosis. —Zoaria branched. Zooecia dimorphic. Large zooecia relatively long, growing parallel to branch axis, and opening only at distal growing tips of branches. Small polymorphs short, budding obliquely from walls of most laterally-disposed zooecia in branch. Apertures of small polymorphs cover side of branch. CERIOPORID CycLosToMEs (Bryozoa): NYE 63 Zooecial walls of large polymorphs granular to indistinctly laminate. Zooecial walls of small dimorphs indistinctly laminate. In zooecial wall, between longitudinally adjacent small polymorphs, laminae form saddle-shaped configurations which arch orally convex longitudinally, but sag transversely, forming aborally convex arches. In zooecial wall, between laterally adjacent small polymorphs, laminae diverge orally at low angles from zooecial boundary zone. Both intermediate and basal diaphragms occur. Taxa included. — The internal characters of C. neocomtensts d’Orbigny, based on an examination of topotype specimens, are con- sistent with the concept of Corymbopora. Fungella, as based on an examination of identified specimens of the type species F. dujardint von Hagenow, is closely similar to Corymbopora in growth habit and, perhaps, other characters. Assess- ment of these apparent similarities must await a more detailed examination of types, or at least topotypes, of F. dujardin. Internal characters of other species assigned to Corymbopora are unknown to me. Discussion. — D’Orbigny (1854, p. 689) recognized dimorphism in Corymbopora: “Chaque branch est terminée par un gros faisceau de cellules verticales . . . La paroi externe des faisceaux est partout criblée en long, de nombreux pores intermédiaires par lignes longi- tudinales”. Gregory (1909, p. 44) noted that there were two sizes of zooecia, but stated that “the sides of the stem are covered by an epizoarial layer, marked by numerous pores, the remnants of the aperture of dead zooecia”. D’Orbigny (1850) and Pergens (1890) synonymized Corym- bopora with Fasciculipora d’Orbigny (1846). D’Orbigny (1854) recognized Corymbopora and Fasciculipora but assigned both to the family Fasciporidae d’Orbigny. Gregory (1909) included Corymbo- pora in the Fascigeridae d’Orbigny in which he also included Fasci- culipora. Borg (1926a) included the genera Domopora and Defrancia in the family Corymboporidae Smitt. Furthermore, Borg (1926a) assigned the Corymboporidae and Fasciculipora to his division 2, the Acamptostega (approximately equivalent to the suborder Tubulo- porina Milne-Edwards, 1838), which are characterized, in part, by single-walled growth of zooecial walls and pseudopores. 64 BULLETIN 291 In all of the above genera, growth habit is similar in that large zooecia grow nearly parallel to the growth axis of the branch throughout their length, and apertures are located only at the distal ends of branches. On the other hand, pseudopores, not small dimorphs, are reported in Fasciculipora (Borg, 1926a, p. 19). The internal characters of Domopora and Defrancia are practically un- known, but illustration of external features of these genera (Borg, 1926a, text-fig. 49, p. 300; text-figs. 79-80, p. 378; text-figs. 83-85, p. 383) do not show the presence of small dimorphs characteristic of C. menardi Michelin, nor were they reported. The wall structure observed in Corymbopora is consistent with Borg’s double-walled concept, but Corymbopora, as presently under- stood, is not easily assignable to Borg’s double-walled groups, the Pachystega (= Horneridae), or the Calyptrostega (= Licheno- poridae). The author has followed Bassler (1953) in discussing Corymbopora with other genera assigned to the Cerioporina (= Heteroporina of Borg), the third group consistent with Borg’s double-walled hypothesis. Thin-sections of specimens assigned to Fungella and Corymbopora neocomiensis Canu and Bassler reveal small dimorphs covering the stem, and other structures suggesting affinity with C. menardi. Detailed study of these species and others is needed before a formal taxonomic action is made, such as possible reassignment of species and erection of a new group equivalent to Borg’s other double-walled divisions. Corymbopora menardi Michelin, 1846 Pl. 10, figs. la-c, 2, 3a-d. 4a-b; Pl. 11, figs. la-d, 2, 3a-b; Pl. 12, figs. 1, 2a-d 1846. Corymbopora menardi Michelin, Iconographie Zoophytologique, descrip- tion par localités et Terrains des Polypiers Fossiles de France et pays environnants, p. 213, pl. 53, fig. 10. 1854. Corymbosa menardi (Michelin), d’Orbigny, Terrain Crétacé Bryozoaires: Paléontologie Francaise Description des Animaux Invertébrés, vol. 5, p. 691, pl. 744, figs. 7-12. Obj. syn. 1909. Corymbopora menardi Michelin, Gregory, Catalogue of fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Cretaceous Bryozoa, vol. 2, p. 45. Type. — A holotype was not designated by Michelin, nor has a lectotype been designated since. Michelin’s primary types are thought to have been originally placed in the Caen Museum, or possibly the Muséum National d’Histoire Naturelle. The museum at CERIOPORID CycLostomes (Bryozoa): NYE 65 Caen was destroyed in the Second World War, and none of Michelin’s specimens of Corymbopora menardi have been found in the col- lections of the Muséum National d’Histoire Naturelle (pers. comm. E. Buge, 1969). Type locality and horizon. — Le Mans (Sarthe), France, Cre- taceous, Cenomanien. Material studied. — Prof. E. Buge kindly made available speci- mens collected by Ferdinand Canu from the topotype locality. The suite of specimens contains a single, nearly complete zoarium, MNHN Canu Coll. 57057-1; and a number of zoarial fragments, MNHN Canu Coll. 57057-2. Thin sections and acetate peels were made of six specimens from MNHN Canu Coll. 57057-2. The original labels indicate that the specimens were collected from “Cenomanien, Le Mans” (Sarthe), France, the type locality. The specimens, MNHN Canu Coll. 57057-2, are labeled as “Fasciculipora (Corym- bosa) menardi d’Orb.” In addition, thin sections and acetate peels were prepared from four specimens in the United States National Museum collection. The specimens, USNM Loc. 2947, were collected from the type locality. Duplicate acetate peels of all specimens sec- tioned are preserved in the United States National Museum collec- tion and the author’s collection. Description. — Mode of growth —Zoaria are small (see Table 8, Zr-Ht, Zr- Wth); branching is dichotomous, producing a delicate, arborescent architecture. Branches have circular cross sections, but expand dis- tally to form capitate distal tips (PI. 7, fig. 1b) with circular to elliptical cross sections (PI. 7, fig. la). Endozone — Zooecia have thin, parallel-sided walls which are locally undulatory but, in general, grow parallel to the growth axis. Walls are homogeneous to subgranular and have thin, dark-colored zooecial linings. Exozone — Remnant, thick-walled growth phases are seen in the capitular growing tips. They are commonly located just proxi- mal or distal to a brood chamber (PI. 10, fig. 3b). The zooecial walls are annularly thickened. The thickenings are roughly symmetrical across adjacent zooecia forming a zone parallel to the zoarial sur- face (PI. 10, fig. 3b). Longitudinally, the zooecial walls have monili- 66 BULLETIN 291 form profiles, symmetrical in thickness across the boundary zone. Thick-walled zones are thin, and commonly have one to four monilar thickenings longitudinally. Small polymorphs — Apertures of small polymorphs are ar- rayed in longitudinal rows on the branch surface. The portion of the zooecial walls between transversely adjacent zooecia forms prominent ridges parallel to the growth axis of the branch (PI. 10, figs. 1b, 2). Small polymorphs have uniformly short, reflexed, conical, or sock- shaped chambers (Pl. 11, figs. 1b, c, e). Interzooidal pores pass through the proximal wall to large polymorphs (PI. 11, fig. le), and also through the walls of longitudinally adjacent zooecia (PI. 11, fig. 1b). Diaphragms — Basal diaphragms were occasionally observed in endozones of large polymorphs. The diaphragms are relatively planar, and show slight oral flexure at the intersection with the zooecial wall. Diaphragms commonly were observed at approximately the same level in several adjacent zooecia. Intermediate diaphragms were observed rarely. The diaphragms flex aborally and merge with the zooecial lining. Brood chambers — Brood chambers are abundant (up to ten were observed in a single zoarium). Abandoned brood chambers were observed only in capitular areas, and are associated with thick- walled zones. Brood chambers are lobate; several lobes branch from a single main lobe, continuous proximally with a single zooecial aperture (PI. 12, fig. 1). Subjacent zooecia are sealed by a thin floor which is structurally continuous with subjacent zooecial walls (PI. 12, figs. 2d, e). Brood chambers observed at the zoarial surface were not roofed over. Abandoned brood chambers were covered by a distal wall bearing interzooidal pores, and growing continuously from the lateral compound wall (PI. 12, figs. 2d, e.) Zooecia budding from the distal wall are commonly thick-walled and have moniliform pro- files °CPI-W2, ‘fies: 2dsre): Discussion. — Athough Michelin’s primary types were not avail- able for examination, specimens were assigned to C. menardi with confidence because: 1) All specimens can be recognized on the basis of external charac- ters, zoarial shape, distally expanded or capitate branches, and CrRIoporiIpD CycLosTomMEs (Bryozoa): NYE 67 the arrangement of the apertures of small polymorphs. These characters are well shown in Michelin’s figure. 2) The species concept, as developed in descriptions or illustrations of D’Orbigny (1854), Gregory (1909), and Canu (1919) has been consistent. 3) Specimens studied by D’Orbigny (1854), Gregory (1909), Canu (1919), and herein were collected from the type locality. Under these circumstances, there are no taxonomic difficulties imposed by the unavailability of the type specimens. Thus, by Art. 7 (a, 1), p. 81, ICZN, 1964, there is no necessity for designating a neotype. Remarks on brood chambers.—In Corymbopora menardi Michelin there is some indication that the brood chamber maintained a connection with a single, presumably fertile, zooecium and hence might be termed a gonozooecium. The use of this term, however, 1im- plies that the whole structure is a zooecial homologue. This is dif- ficult to apply because the wall which bounds the brood chamber is compound, and shared with subjacent and lateral zooecia. The de- positing epithelium is probably better considered as a zoarial rather than a zooecial tissue. For this reason, the more general term, brood chamber, is here retained. In C. menardi, the brood chamber was never roofed over by a porous, diaphragm-like structure typical of the brood chambers in Ceriocava corymbosa (Lamouroux), Heteropora cryptopora (Gold- fuss) and Parleiosoecia jacksonica Canu and Bassler. The brood chambers in C. menardi were apparently covered by calcareous tis- sue only after abandonment. The covering structure is similar in appearance to the adjacent zooecial wall rather than the basal, layer- like structure seen in other genera. Genus COSCINOECIA Canu and Lecointre, 1934 Type species: Coscinoecia radiata Canu and Lecointre, 1934, p. 21, by original designation and monotypy. 1934. Coscinoecia Canu, and Lecointre, Les Bryozoaires Cyclostomes Des Faluns de Touraine et d’Anjou, Soc. Géol. France, Mém., vol. 9, No. 4, p. 198. 1953. Coscinoecia Canu and Lecointre, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G70. 1957. Coscinoecia Canu and Lecointre, Buge, Mus. Nat. d’Hist. Nat., Mém., ser C, Sciences de la Terre, vol. 6, p. 121. BULLETIN 291 68 SI9JIWINI|[TU UT, T £ Ser $21 0 ‘S US'D9Z/UD-7d PZ i £ Scr Sct 163 £00° OO $00° cc" YL-IM9ZPO DGEINNGES§D 21027 I £ Sct Sct | co fT OT Ve UGXI-4S8§9-4D9Z T £ Scr rat 61 c0" aS +0° LY UOXINN-USS8)0-YO9Z T £ Scr Sct LT 20° tT 80° 0c UOXxN-8889-4D°Z auozopuy - sydiowAjog aS1eq - [e102007 9 ‘TI PM-1Z 9 “€T IH-AZ [Bl1e0Z7 2pop29dg = IZN-oOOZN N ‘AO «8 #X «WO PeETENS NITHHOIN IGYPNAW PYOdOIWANOD AO SLNAWAAASVAW AO AUVWIWAS IVOLLSLLV.LS 8 HIAVL 69 CERIOPORID CycLosToMEs (Bryozoa): Nye ‘I-LS0L5 ‘OD NUL NHNW :2dAjodoy, *(L) ‘(9) £(Z) “(S) (4) ‘(h) § (4) (1) 225025 ‘1190 nUeD NHN OMe (A eu(S a ( “CSTE CS) C8) ce) sens BO) Cb)! on9)) he TOS) iS, CS) a 0Se( 9) Z-LS0L$ ‘1190 nUeED NHNW £) @-LS0LS “19D nue NHN £) 2-LS0L$ ‘1l9D0 Due) NHN adOO NAWIOUdS OL ATH :sadAjodoy, * :sadAjodo 7, :sadAjodo [, :sadAjodo T, ° :sadAjodo [, aN HW tH S SIDJIUNT| [IU UT y S + 0Z + ¢ a 3° Zc Wd-yold + Z 81 L £0° 09° St 06° qT M-YOld Jaqueyy) poolig Z z 9T 91 ££ z0° $0" £0" L0° ALLY L-LM9ZPO 4 Z rai ra We 10° +0" z0° $0" 8uTY,L-LM9ZPO C g cl rat ZI 10° $0 +0" 90° UdAI L-USsD-YDIZ Z zZ rat ra 61 10° $0" +0" L0° uqsuyT-ugs)-YyD9Z sydiowdjog |[BWS - [e109007 apo ‘90dg = IZN-COOZN N 178) *S 4X «wo JayoereyS NITHHOIW IGYPNAW FAYOMOIWAUYO)D AO SLNANAYNSVAWN AO AUYVNWWAS TVOILSILV.LS 70 BULLETIN 291 Tentative diagnosis.— Zoaria branched; branches have well- differentiated coaxial endozones and exozones. Zooecia dimorphic. Large polymorphs occur in intermonticular areas, small polymorphs in intermonticular and monticular areas. In exozone, zooecial walls distinctly laminated. Laminae broad- ly arched across zooecial boundary zone. Laminate tissue sometimes alternates with granular tissue which forms discontinuous rounded masses in outer cortex. Basal diaphragms numerous and closely spaced in endozone and zone of flexure. Intermediate diaphragms occur in intrazoarial overgrowths. Taxa included. — Monotypic for the type species, C. radiata. Discusston. — Coscinoecia is a large bryozoan, distinctive both externally and internally. Bassler (1953, p. G70) assigned Cosctnoe- cia to the family Tetrocycloeciidae Canu and noted in the diagnosis that Coscinoecia is “like Tretocycloecia [sic] but oeciostome larger than zooecial apertures”. Coscinoecia is similar to Tetrocycloecia, Leiosoecia, and Ceriocava in its coaxially ramose growth habit. Coscinoecia, however, differs from all three genera in several charac- teristics, such as the structure and appearance of the zooecial walls, the arrangement of dimorphs, and the occurrence of diaphragms. The gross morphologic structures are consistent with placement in the Cerioporina, but the differences in morphology are too great, as presently understood, to indicate close affinities with any genus studied here. Coscinoecia radiata Canu and Lecointre, 1934 Pl. 13, figs. la-h; Pl. 14, figs. la-h; Pl. 15, figs. la-i 1934. Coscinoecia radiata Canu, and Lecointre, Soc. Géol. France, Mém., vol. 9, No. 4, p. 198, pl. 40, figs. 1-7. 1957. Coscinoecia radiata Canu and Lecointre, Buge, Mus. Nat. d’Hist. Nat., Mem., ser. C, Sciences de la Terre, vol. 6, p. 122, No fig. Type.—Canu and Lecointre (1934) noted that they had examined several specimens from seven localities in the Helvetien of southern France. One specimen from the primary type suite has been located. This specimen, MNHN Canu Coll. 58872, is the large, well-preserved specimen figured by Canu and Lecointre (1934, pl. 40, figs. 1-4) and is here designated as the lectotype. Thin-sections were figured by Canu and Lecointre (pl. 40, figs. 5-7); the specimen CERIOPORID CycLosToMEs (Bryozoa): NYE 71 or specimens sectioned were not identified. The thin-sections were not located by the author. Although Buge (1957, p. 122) referred to MNHN Canu Coll. 58872 as the holotype, there is no such desig- nation in the primary source. Type locality and horizon. — Canu and Lecointre (1934, p. 198) cited the following localities in France for the occurrence of C. radiata: Faluns (Helvetien) Indre-et-Loire: Saint Epain, Ferriére-l’Arcon Marn-et-Loire: le Haguineau, Doué-la-Fontaine, Montlouet Ille-et-Vilaine: environs de Rennes Burdigalien supérieur Piémont: Croce Barton The label with the lectotype lists the word Doué; therefore, the type locality and horizon are restricted to the Faluns (Helvetien) [Miocene], Doué-la-Fontaine, Marn-et-Loire [France]. Material studied. — The lectotype was kindly loaned to the author by Dr. E. Buge, Muséum National d’Histoire Naturelle. Five thin-sections and 13 acetate peel replicas were made from the lecto- type; most of the original zoarial fragment and three remnants re- main after sectioning. Duplicate peels are in the United States National Museum and the author’s collection. Description of the lectotype. — This description is based solely on the lectotype, the only specimen of C. radiata known to be avail- able for study. This description includes assessment of some non- genetic variation within this colony. No assessment of genetic or other interzoarial variation within C. radiata is implied in this description. Mode of growth. — The zoarium is robust (Table 9, Zr-Ht, Zr- Wth, Br-CsSn-MxDn); branches are roughly cylindrical. Most of the surface of the primary branch is encrusted by an intrazoarial overgrowth (PI. 13, figs. le, f). Large polymorphs occur in intermonticular areas, and are ar- ranged in rows disposed radially about the monticular areas (PI. 13, figs. 1b, c). Small polymorphs occur in intermonticular areas, commonly in rows between the large polymorphs (Pl. 13, figs. 1b, c), and also in monticular areas (PI. 13, figs. 1b, c, h; Pl. 14, fig. 1d). Less regular arrangements occur locally (PI. 13, fig. 1h). 72 BULLETIN 291 Endozone. — Zooecial walls are thin, symmetrically to subsym- metrically thickened across the boundary zone, and have submonili- form profiles. Zooecial chambers commonly have subelliptical, sub- polygonal cross sections. Zooecial walls are generally granular, some- times indistinctly laminate, and a thin zooecial lining is usually present. Basal diaphragms are commonly thin and planar to slightly convex aborally. At the juncture with the zoecial wall, diaphragms flex orally and merge with the zooecial lining (PI. 13, fig. 1d; PI. 15, figs. lib, 1): Exozone. — The zooecial walls of adjacent large polymorphs are symmetrically to subsymmetrically thickened across the zooecial boundary zone and have moniliform profiles, largely because of the occurrence of numerous, widely-flexed, interzooidal pores (PI. 13, fig. 1g). Monilar profiles commonly are elliptical, ovate or fusiform, but show less variation in thickness longitudinally than small poly- morphs, and appear nearly parallel-sided (PI. 13, figs. le, g). Zooecial chambers commonly have subelliptical cross sections but show relatively large variation in this respect (Table 10). Cross sections are often somewhat irregular because of crescentic inflec- tions in the zooecial wall (PI. 14, fig. lh). Zooecial walls of small polymorphs are commonly thickened an- nularly. Back-to-back adjacent small polymorphs are thickened sym- metrically across zooecial boundary zones (PI. 13, figs. le, g; Pl. 14, figs. la, b, e), but are commonly thickened asymmetrically across zooecial boundary zones when adjacent to large polymorphs (PI. 13, fig: Ig; Pl. 14; figs: 1b, f; Pl. 15, fig. 1g). Monilar>profilesof adjacent small polymorphs are circular, elliptical, clavate, inverse- pyriform or alate. In monticular areas, cross sections of small poly- morph chambers are subelliptical to subcircular (Pl. 14, fig. 1d). In intermonticular areas, cross sections of small polymorph chambers are commonly more elliptical and less regular. Small, blunt mural spines were often observed in both large and small polymorphs (PI. 14, figs. 1d, g). Diaphragms. — Basal diaphragms were observed only in the endozone and zone of zooecial bending. The diaphragms are numerous, closely spaced (Table 9, Bsl-Intvl), and generally thin (Table 9, BsID-Th). The diaphragms are planar to slightly convex 1S Crrroporip CycLosToMEs (Bryozoa): NYE S19JIUIT|[IU UT» I I OOT OOT 0 £ ugsp9Z/4D-1dPZ I if Oot Oot $$ 620° c$0° O10" oot YL-LM9ZPO UCGXWN-USS0-YOOZ I T oor Oot Le 9¢° al OT ge UCXIN-USSD-YO2Z I i Oot Oot Le +0" or £0° LV UCGXWN-YUSSO-4O9Z I I OOT OOT ce +0" cr $0° +2 WAXW-USSD-YO9Z auozoxg - sydiow jog |[Y - [B19900Z7 I I c $ IT udxW-USs)-14 I I I OF WM-41Z i I I 0s HAZ [Bl1e0Z apop 2edgs_ IZN (OZN N “AO #S «X PRA O) ALLNIOOAI GNV ONVO PLFIGFVY FIDFONIISOD AO SLNANAYASVAW AO AUVNWNAS TVOILSILV.LS 6 ATaVL BULLETIN 291 74 SIDJOWIN] [IW UT y T I LS LS 91 Z0° tT 80° 8t° UQxXIW-4S§89-49Z sydiows]og adie - [e109007 T T $c Sc 0 c Ugs99Z/UD-1dPZ T T $c $c $é 400° Z10° $00° 610° YL-LM9PZPO UG*WN-4S80-YO9Z T I $c $Z IT ev cl OT Suk UCXIN-4SsSD-YOIZ T T Sz $c $c £0° ae +0 hs UGXWN-USSD-YO2Z T T $c $c <4 +0° st +0" st UdGXxW-USsSD-YO2Z auozopuy - sydiowdjog [[V - [2199007 I I oor 00r 0 9 ugs2Z/UD-dS9Z (‘uod) auozoxg - sydiowsjog [[V - [8199007 apo ‘oads IZN 9ZN N AGO aS #X «WO sapBIVY) AALNIOOAT GNV ANVO PLVIGFPY FIDFONIDSOD JO SLNAWNAYNSVAW AO AUVNWAS TVOILSILVLS 75 CERIOPpoRID CycLosToMEs (Bryozoa): Nye S1O}IUIT] [IU UT» UQXIWN-4S80-4YO9Z T T ch ch <4 be tT OT 6¢ UGXW-4S8)D-4D9Z T I cb ch 97 Z0° 90° £0" Il’ UQXNWN-US8)-YO9XZ T T ch cr 97 ZO" 60° $0° o) UQXI-USs0-YO2Z Iv[nojuowsajuy - sydiowAjog [[Bus - [8199007 I I LS LS 0 9 9Z /WD-4$9Z T T Ls LS 0 £ US8)9Z/UD-1d PZ T i LS LS tr 020° $+0° 010° 260° WL-TMPZPO UGXINN=9S8O=4D9Z I I LS Es 02 o cr OT oC BOXIN-USSOSN02Z I T LS LS 02 zo" cr 90° LV UQXWN-"S8D-4D9Z (‘uod) sydiowAjog asivyq - [8199007 apoD ‘oad TZN %ZN N AO aS aX «WO taeIBY) GULNIOOYT GNV ONVO PLIVIGFY PIDZONIDSOD AO SLNANAYNSVAW JO AUVWAAS TVOILSILV.LS BULLETIN 291 76 “ZL88S “I19D NUE NHNIW edAjoq097] “1 adoo NAWIOddS OL AUM ‘BI19900Z Jou {siaquIBYyD poo1q Fo Jaquinnt SId}JIWN [IW UT, T I +£ cl Lt 90° se 67° cs Wd-YOold T T +1 I TT WM-YOl"d T T +1 I tT WT-Yord Jaqueyg pooig T I Sc $¢ or Or 9e° $$" 10]-CIsa auozopug - wseiydeiq I I L L £9 Z00° +00° z00° 800° WL-CIsa I T 9 9 9S Z00° +00° c00° 900° YL Gs] auo0zoxy - wdseiydeiqg T if $ $ OT 800° 80° 18° Tor YL-TLM9ZPO T T 6 6£ +c z0° L0° UGXINN-YSS80-YO9Z T if 6£ 6£ $c Z0° 60° UQXIN-USSD-YOIZ Ie[noyuoyy - sydiow jog |[BWs - [e199007 I I cb cb 0 $ 9Z/UD-4$9Z I I ch ch 0 c Ugs)9Z /UD-1dPZ T T cb ch 8S 9£0° 790° 0TO" oot YL-TAAPZPO (‘uod) Iev[NoyUOUIazUY - sydiow dog [[BUISg - [2199007 apoD ‘eds IZN %ZN N AGO «S aX «WO JayoB1eYy) ALLNIOOAUT GNV ANVO PLVIGPY FIDAONIDSOD AO SLNAWAYANASVAW AO AYVWWAS TVOILSILV.LS Crerroporip CycLostomeEs (Bryozoa): Nye Vi aborally, and flex orally to form abutments before merging obscurely with the zooecial lining (PI. 15, figs. 1b, i). Most zooecia in the intrazoarial overgrowth have a single, thin (Table 9, IntD-Th) intermediate diaphragm at approximately .2 mm aboral to the skeletal aperture (PI. 15, fig. la). The diaphragms are generally planar, and they flex aborally to form a short abut- ment which adjoins, but does not merge with, the zooecial wall. Brood chambers. — Brood chambers are large (Table 9, BrCh- Lth, BrCh-Wth, BrCh-Dth) zoarial cavities with lensoid to rec- tangular profiles (PI. 15, figs. 1c, f) and sublobate cross sections (PI. 15, fig. le). The floor is incomplete (PI. 15, fig. 1c), and subjacent zooecia show various degrees of closure, from fully open to completely sealed. Closure may be made both by lateral growth of the apertural portion of the zooecial wall, and by emplacement of a nonperforate intermediate diaphragm (PI. 15, fig. 1h). The roof is apparently nonporous (Pl. 15, fig. 1c). Small polymorphs bud from a thin basal layer which encrusts the roof (PI. 15, fig. 1d) forming monti- cular areas. The apertures of these small polymorphs are commonly elliptical to slitlike, in contrast to the more rounded openings of normal monticular areas. Encrusted brood chambers commonly open to the surface by means of a single, large, subcircular to elliptical opening (PI. 13, figs. 1b, c). TABLE 10 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN THE LECTOTYPE OF COSCINOECIA RADIATA CANU AND LECOINTRE Cire. — Ellip. Ov. Pyr. Polyg. Triang. Irreg. Exozone Regular 1 28 ai 3 1 Sub 3 31 al 1 Irregular 3 4 2 5 Endozone Regular 2 1 1 Sub 1 10 3 5 1 Irregular 1 78 BULLETIN 291 751A 75 B ~~ ~> oO 12) = SC oO 250 250 lox S: wo o 25 25 mm 4a of <5) 1.0 2.0 ZcCh-CsSn-NMxDn ZcCh-CsSn-Mx Dn ZcCh-CsSn-NMxDn 754C ~> a 100 = © 5 90 SSS o 5 w ne > - 7s 350 25 au Ss & 25 =} oO _ mm 4 a2 We 2 bub CdZcW!-Th ZdPr-Cn/ZcCsSn Text-figure 8 A-D. Histograms and cumulative curve from the lectotype of Coscinoecia radiata Canu and Lecointre. A. Normal to maximum cross- sectional dimension of a zooecial chamber. B. Ratio of the maximum cross- sectional dimension of a zooecial chamber to the normal to maximum cross- sectional dimension of a zooecial chamber. C. Compound zooecial wall thick- ness. D. Count of interzooidal pores per zooecial cross section. Cerioporip CycLostomes (Bryozoa): NYE 19 Remarks concermng morphology. — Growth habit — The erect, robust growth habit of C. radiata provided a suitable substrate for other bryozoan colonies. Several small cheilostome colonies and one lichenoporid colony were ob- served on the zoarial surface. Intrazoarial overgrowth could be interpreted, in the situation figured in Plate 13, figures le, f, as a mechanism of defense because the overgrowth encrusts a relatively large cheilostome colony which had submerged part of the main stem. Polymorphism — The starlike pattern engendered by the ar- rangement of dimorphic zooecia is a distinctive external character. In tangential sections the pattern is not always so readily visible and requires comment. Remarks on this subject must be considered tentative, however, because of limited material. When viewing the zoarial surface, the eye readily makes the distinction between large and small polymorphs. Large polymorphs have relatively constant zooecial apertures in the exozone, and one sees a large, black zooecial opening (PI. 13, figs. 1b, c). On the other hand, small polymorphs have much more variable cross-sectional areas because of pronouonced annular thickening of the walls. When the zoarial surface is viewed, the eye “sees” the smallest cross section of small polymorphs as a black void; indeed, the largest dimension is nearly ignored unless an effort is made to see it (note light-colored areas around black voids at top, fig. 1c in PI. 13). In sections, zooecia are cut at all levels. If the section intersects small polymorphs where walls are thickest, the polymorphs are easily distinguishable. Conversely, where walls are thin, the areal dimensions approach, and perhaps overlap, the large polymorphs (see Text-fig. 8A). Thus, the regular pattern, so distinct at the zoarial surface, is obscure in thin-section. Brood chambers —In comparison to brood chambers in Certo- cava, Heteropora and Parleiosoecia, Coscinoecia has brood chambers with nonporous roofs and incompletely calcified floors. Another difference is that a single, large connection with the surface is retained well after the calcareous brood chamber roof was deposited, suggesting that larvae may have been contained, or brooded, after the chamber was roofed over and submerged beneath new growth. 80 BuLtetTIn 291 Genus DIPLOCAVA Canu and Bassler, 1926 Type species: Diplocava incondita Canu and Bassler, 1926, p. 71, by original designation. 1926. Diplocava Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 67, p. 71. 1953. Diplocava Canu and Bassler, Bassler, Treatise on Invertebrate Paleon- tology, Part G, Bryozoa, p. G71. Tentative diagnosits.— Zoaria massive to branching. Growth habit variable, with complete gradation between two end members. In one growth habit, most zooecia relatively short and growing radially to produce lenticular intrazoarial overgrowths. In second habit, axial zooecia longer, initially growing nearly parallel to major growth axis before bending towards zoarial surface, and producing coaxial appearance; but, zooecial walls commonly not showing typical endozonal appearance in axial region. In exozone, zooecial walls laminate. Laminae diverge orally from generally dark-colored, narrow boundary zone; then arch convex orally and recurve aborally. Cortex laminae abut thin zooecial lining at low angle. Walls commonly have integrate appearance in tangential section. Simple external walls common. Intermediate and basal dia- phragms occur. Taxa included. — Only the type species was studied. Internal characters of other species assigned to Diplocava are unknown to me. Discussion. — Diplocava was originally defined as “Ceriocavidae with dimorphic zooecia” (Canu and Bassler, 1926, p. 71). The dimorphs were described by Bassler (1953, p. G71) as“... large open ones separated by a zone of small ones with facets.” Examina- tion of thin-sections reveals, however, that zooecia are monomorphic. Another interpretation of the divergent zooecial appearance seen in Diplocava is given below. The emplacement of simple external walls apparently represents a late phase in zooecial ontogeny. The portions of the compound zooecial wall adjacent to the external simple wall are consistent in having thick walls with recurved laminae, in having a well-defined boundary zone, and in having nearly parallel-sided profiles, charac- teristics not seen in more aboral portions of the zooecial wall. In addition, zooecia of each type occur in groups. Zooecia with large apertures encrust the zooecia sealed by external walls (“small CrerroporIp CycLtostomes (Bryozoa): NY 81 ones with facets”) (PI. 16, fig. 1b). The present interpretation sug- gests that the “large zooecia” were in a stage of active, orally-directed growth, while the “small zooecia” had ceased active, orally-directed growth of the zooecial wall and had secreted simple external walls at the skeletal aperture. The Ceriocavidae were defined by Canu and Bassler (1922, p- 89) on the appearance of the brood chamber which they described as “long, transverse, convex, regular, symmetrical vesicle with special walls”. Although Canu and Bassler (1926, p. 71) noted in their remarks on the type species that the brood chamber was unknown in Diplocava, they described (on the same page) a “star-shaped” brood chamber from a “cotype” of the type species. This specimen is con- sidered here to be assignable to another, perhaps unnamed, genus. Remarks concerning its morphology, 1.e., brood chamber, are not applicable to the concept of Diplocava. The assignment of Diplocava to the Ceriocavidae Canu and Bassler, based on the brood chamber alone, is not justified. Presently, lack of knowledge of changes in skeletal structure through time in cyclostomes makes phylogenetic inferences speculative; however, in- ferences of close relationship between Ceriocava and Diplocava can- not be supported by morphologic evidence as understood here. The respective type species differ in growth habit, wall structure, and occurrence of diaphragms. They differ, as well, in other characters tentatively considered significant at the specific level, e.g., profiles of zooecial walls, and outlines of zooecial chambers. Simple external walls have been recognized in Haploecia. Diplo- cava differs from Haploecia in mode of growth and wall structure. Diplocava and Reptonodicava both have recurved laminae in the outer exozone. Reptonodicava differs from Dzplocava in mode of growth and occurrence of diaphragms. Diplocava incondita Canu and Bassler, 1926 PIVIGs figs la-hs eles: figs. 1-4, 5a-c, 6a, b; Pl. 18, figs. 1-3; Pl. 19, figs. 1-3 1926. Diplocava incondita Canu and Bassler, U.S. Nat. Mus., Proc., vol. 67, pp. 71-73, pl. 10, figs. 5 (specimen in lower right corner of figure), 6. Type. — USNM 69925-2 figured by Canu and Bassler (1926, pl. 10, figs. 5 — specimen in lower right corner of photograph — and 6) is designated as the lectotype. Five unnumbered thin-sections labeled type are paralectotypes. 82 BuLLETIN 291 Type locality and horizon — Lower Cretaceous (Valangian), Sainte Croix (Vaud), Switzerland. Material studied. — Five thin-sections and five acetate peels on one slide were made from the lectotype. In addition, 22 thin-sections and 24 acetate peels were made from 10 specimens collected from the type locality (USNM Loc. 2404) and identified as Diplocava 1n- condita by Bassler. Duplicate acetate peels are preserved in the bryozoan collection of the National Museum of Natural History and in the author’s collection. Description. — Mode of growth — Zoaria are small (maximum observed length is about 11 mm, diameter about 5 mm), and are either irregularly domal to digitate masses or are branched with subcylindrical branches. The irregular zoarial appearance (PI. 16, fig. 1a) is caused by the irregular repetition of both encrusting and coaxial modes, and by the variation in position and direction of the major growth axis from one overgrowth unit to another. Intrazoarial overgrowths sometimes cover most of the surface of subjacent growth units (Pl. 17, fig. 3), or are restricted to the surface peripheral to the continuously-growing zooecia of the axial region (PI. 16, fig. 1d; pl. 17, fig. 5a). Basal layers are generally thick (Pl. 16, fig. 1g; Pl. 17, figs. 5b, c, 6a, b). Proximal parts of overgrowths sometimes display thin, recumbent, zooecial walls inter- preted as endozones (PI. 16, fig. 1h, on left); but commonly, zooecial walls are essentially thick-walled throughout. Exozone — The zooecial walls are symmetrically thickened across the boundary zone. The walls generally show a regular in- crease in thickness orally. Sometimes, however, the walls show a slight thinning in medial portions (PI. 17, fig. 3). In the inner exo- zone, thinning of the zooecial wall near interzooidal pores commonly produces moniliform profiles (PI. 17, figs. 1, 5c). In the outer exo- zone, interzooidal pores are almost cylindrical with little flare (PI. 17, fig. 6b; Pl. 19, fig. 1), resulting in nearly parallel-sided zooecial walls. Slight pustulose thickenings are sometimes seen in the outer exozone (Pl. 16, fig. lg). Cross sections of zooecial chambers generally are smoothly rounded and subcircular to elliptieal (PI. 19, fies li) 83 CERIOPORID CycLosToMEs (Bryozoa): NYE SId}IWII[ [IW UT, 4 b ff fe Te TT0° 9£0° 9T0° 190° YL-MdS eA [eusaxq ajduis £ $ 8Er 8ér 0 BS IQUIN-1dPZ c $ 6ET 6ET SE 970" 890° STO 9bT YL-LM9ZPO UQXINN-US§8D-YO9Z c $ 6£T 6ET IT v raat OT LT UQXW-USsD-YO2%Z é $ 6eT 6ET cc +0’ LY +0° 9o° UQXNN-YS8)-YO9IZ c $ 6£T 61 02 +0° 0c so 8c UdxXW-USsD-YOXZ aU0ZOXY - [2199007 T $ € 5S UQXI-USs0-1¢4 [BI1B0Z apop29dgs =IZN ZN N ‘AO «8 «X «WO qajovreyO YAISSVA GNV ONVO PLIGNODONI FAFIOTdIC AO SLNAWAYNSVAW AO AUVNWIANAS TVOILLSILV.LS Il ATa#VL BULLETIN 291 ‘(p) 8-bObe ‘(£) L-bObZ ‘(T) I--0bZ “20T WNSA ‘42a]ssegq Aq palfznuapr *(OL) O1-FO+Z ‘(O1) 8-tOFZ ‘(0Z) I-b04% “997 WNSQ ‘4alsseg Aq palfznuapr “8-bObC °8 ‘8-t0bZ pue adAjoaqT ‘7 ; (9) OL-b0+z ‘AW[BIO] adA} ay} WOLF pajzoa[[od suauIoads aay T *9 ‘AY[BIO] ad} 2Y} WOIZ pazoa]joo suautoads aaiyy, “¢ "(9) 8-40b2 ‘(9) 9-b0bZ ‘(Z) T-40tZ ‘207 IWWNSY :42[sseg Aq palynuspr ‘Aypeoo, addy ay} Wolf pajda[[oo suauaads aaryi ! (+1) 7-$7669 IWNSQ :9dA}01997 “+ ‘(LE) €-bOte }daoxa 7H se auieg ‘¢ "(9) S-bOtZ ‘(Tb) t-b042 ‘(8E) €-bObz ‘(LZ) I-b0+Z WNSQ :49[sseg ‘S ‘y Aq palguuapr pue Ajljeoo] adA} ay} WOIF pajda[jood suauoads ANOF ! (OE) 7-$7669 INNSA :adAjojaT °Z "2-$7669 INNS :adAjo}097 *T ad0O0 NYWIOddS OL ATM SI9}JIUWIT[[TW UT, L 4 > ¥ 200° £10" WL-dIsa 8 I I I 800° een Ae Elon LA | suseiydeiq $ £ 8 Ot cc c00° TT0° 800° 120° UCXIN-USSD-1dd-dPpsd-MaAS 9 tI tI 6c cc0" cL0° 950° £60° UG*AN-USSO- Mas HEA [BUta}xq a[dwis apop9edg = IZN 9ZN N AD *S *X «WO ESE £D) UAISSVA UNV ONVO PLIGNOONI FAVOOTdIC AO SLNAWNAYNASVANW AO AYVNNAS TVOLLSILV LS Cer1oporIp CycLostomes (Bryozoa): NYE 85 75,4 75 B > ~> oO oO (= (‘= 250 e50 o o 2 : 25 2S mm 4a v2. ae 1.0 2.0 ZcCh-CsSn-NMxDn ZcCh-CsSn-Mx Dn ZcCh-CsSn-NMxDn 754C ~> & 100 = o 5 90 x75 oT ® : - 21250 25 ~ =) P45) = | oO mm A Pe ul PA Sie Sy GS CdZcW!-Th ZdPr-Cn/ZcCsSn Text-figure 9 A-D. Histograms and cumulative curve from the lectoiype and four topotypes of Diflocava incondita Canu and Bassler. A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness. D. Count of interzoidal pores per zooecial cross section. 86 BULLETIN 291 Simple External Walls — Simple external walls (PI. 16, figs. 1d, o; Pl. 17, figs. 5b, c,.6a, b;4Pl. 18, fies: 1, 2,°3; BE 19g yop eveene observed in most zooecia subjacent to intrazoarial overgrowths. The walls are thick, and apertures generally located in the disto-central or central part of the diaphragm. A deposit of laminated calcareous tissue, continuous with the zooecial lining, lines the aboral surface of these diaphragms sealed by overgrowths (PI. 16, fig. 1g; Pl. 17, fig. 6a; PI. 18, figs. 1, 2) and extends orally to line the peristome. Diaphragms — Intermediate diaphragms occur rarely and, when seen, are just aboral to the aperture (PI. 19, fig. 2). The diaphragms are thick, laminate, and have prominent aborally flexed abutments. Basal diaphragms occur rarely. The diaphragms are thin and nearly planar, and flex orally at the juncture with the zooecial wall to merge with the zooecial lining (PI. 16, fig. 1g; Pl. 18, fig. 2). Discussion. — Three of the four original specimens figured by Canu and Bassler (1926, pl. 10, fig. 5) were attached to a small card bearing the label “Diplocava incondita, Cotypes, USNM 69925”. Two of the specimens were thin-sectioned. USNM 69925-2 was judged to be conspecific with unfigured (?) thin-sections prepared by Bassler and labeled as types by him. In addition, USNM 69925-2 was judged to be conspecific with sectioned non-type specimens from the type locality identified by Bassler as D. incondita. The second specimen sectioned, USNM 69925-1, is a thin-walled encrusting bryozoan referable to a different, probably unnamed, TABLE 12 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN DIPLOCAVA INCONDITA CANU AND BASSLER Gao Ellip. Ov. Pyr. Polyg. Triang. Irreg. Endozone Regular 2 39 6 Sub 10 66 6 2 Irregular a 3 1 Exozone Regular 6 2 Sub 5 + Irregular CERIoporID CycLosToMEs (Bryozoa): NYE 87 genus. This was the only specimen of Canu and Bassler’s original suite in which a brood chamber was observed. Thus, Canu and Bassler’s remarks concerning the occurrence and morphology of brood chambers are not applicable to the concept of D. incondita. Genus DITAXIA von Hagenow, 1851 Type species: Ceriopora anomalopora Goldfuss, 1826, by sub- sequent designation, D’Orbigny (1854, p. 952). 1826. Pars Ceriopora Goldfuss, Petrefacta Germaniae, vol. 1, p. 33. 1830. Pars Heteropora Blainville, Zoophytes, Dictionnaire de Science Naturelles, vol. 60, p. 382. 1834. Pars Heteropora Blainville, Blainville, Manuel d’Actinologie ou de Zoophytologie, p. 417. 1851. Ditaxia von Hagenow, Die Bryozoen der Maastrichter Kreidebildung, Cassel, p. 49. 1854. Ditaxia von Hagenow, D’Orbigny, Terrain Crétacé Bryozoaires, Palé- ontologie Francaise Description des Animaux Invertébrés, vol. 5, p. 953. 1881. Polytaxia Hamm, Inaugural-Dissertation zur Erlangung der Doctorwurde Von der Philosophischen Facultat der Friedrich-Wilhelms-Universitat zu Berlin, p. 41. 1899. Ditaxia von Hagenow, Gregory, Catalogue of fossil Bryozoa in Depart- ment of Geology, British Museum (Natural History), The Cretaceous Bryozoa, vol. 1, p. 406. 1922. Ditaxia von Hagenow, Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 61, p. 101. 1953. Ditaxia von Hagenow, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G72. Tentative diagnosis.—Zoaria_ encrusting and_ branching. Branches bifoliate and commonly frondose, less commonly sub- cylindrical. Endozones and exozones strongly differentiated and intergrade in narrow zone of zooecial flexure. Zooecia dimorphic. Large dimorphs occurring in intermonticular areas; small dimorphs occurring in intermonticular and monticular areas. In exozone, large dimorphs have lunaria-like structures com- posed of subgranular calcite. Lunaria-like structure becomes obscure near aperture, merging with indistinctly laminate tissue. Thin zooecial lining commonly present. Small polymorphs have discontinuous patches of homogeneous to subgranular calcite in cortex, alternating with indistinctly lami- nate tissue. Laminae crenulate, broadly curved convex orally, and sometimes continuous across zooecial boundary zone. Intermediate diaphragms occur rarely. Taxa included. — Only the type species, D. anomalopora; in- 88 BULLETIN 291 ternal characters of other species assigned to Ditaxia are presently unknown to the author. Discussion.—In the large zooecia of Ditaxia, most of the cortex of the proximal wall is composed of calcareous tissue which is distinctly different from calcareous tissue in the remainder of the wall. This light-colored, homogeneous tissue is called a lunaria-like structure here, and is inferred to have been originally granular be- cause remnants of originally laminate tissue are preserved elsewhere in the zooecial walls. Recently, studies of lunarial microstructure were made in some Paleozoic cystoporates by Utgaard (1968a, b), and in post-Paleozoic lichenoporids by Brood (1970b). Unfortunate- ly, comparison of lunaria in these taxa to the lunaria-like structure in Ditaxia cannot be made because microstructure is poorly pre- served in the specimens of Ditaxia which are available for study. The recognition of a lunaria-like structure poses some questions concerning continued assignment of Ditaxia to the cerioporids. On the basis of mode of growth, wall structure, and diaphragms, Ditaxia is not readily assignable to other taxa as presently understood, e.g., lichenoporids and is provisionally retained in the cerioporids. Ditaxia anomalopora (Goldfuss), 1826 Pl. 20, figs. la-e; Pl. 21, figs. 1, 2a-c; Pl. 22, figs. 1, 2a-c 1826. Ceriopora anomalopora Goldfuss, Petrefacta Germaniae, vol. 1, p. 33, pl. 10, figs. Se, d. 1830. Heteropora anomalopora (Goldfuss), Blainville, Zoophytes, Dictionnaire de Science Naturelles, vol. 60, p. 382. 1834. Heteropora anomalopora (Goldfuss), Blainville, Manuel d’Actinologie ou de Zoophytologie, p. 417. 1851. Ditaxia anomalopora (Goldfuss), Von Hagenow, Die Bryozoen der Maastrichter Kreidebildung, Cassel, p. 49, pl. 4, fig. 9c. 1881. Polytaxia anomalopora (Goldfuss), Hamm, Inaugural-Dissertation zur Erlangung der Doctorwurde Von der Philosophischen Facultat der Friedrich-Wilhelms-Universitat zu Berlin, p. 41. 1899. Ditaxia anomalopora (Goldfuss), Gregory, Catalogue of Fossil Bryozoa in Department of Geology, British Museum (Natural History), The Cretaceous Bryozoa, vol. 1, p. 406. 1922. Ditaxia anomalopora (Goldfuss), Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 61, p. 101. Type. — Specimen UB 119 is designated as the lectotype. The lectotype was figured by Goldfuss (1826, pl. 10, figs. 5c, d) and by Von Hagenow (1851, pl. 4, fig. 9c). Type locality and horizon. — Goldfuss (1826, p. 33) cited the Crerroporip CycLostomes (Bryozoa): NYE 89 locality as “Petersberge bei Maastricht”. Rocks exposed at this locality are Maastrichtian in age. Therefore, the horizon is assumed to be Cretaceous, Maestrichtian. Material studied. — Four thin-sections and four acetate peel replicas were made from the lectotype UB 119; most of the lectotype remained after sectioning. In addition, eight topotypes in the collec- tions of the National Museum of Natural History were thin-sec- tioned: four specimens labeled “USNM Loc. 2405, Up. Cret.-Maastr., Geulem, Maastricht, Netherlands”; and three specimens labeled “USNM Loc. 2965A, Cret.-Maastr., Maastricht, Netherlands”. Duplicate acetate peels of specimens thin-sectioned are in the Bonn collection and the National Museum of Natural History collection. The author has duplicate peels of the lectotype and the specimens from USNM Loc. 2405. Description. — Median layer — The median layer is thick (Table 13, MdnLyr- Th) and laminate. In cross section, the laminae form steep V- shaped configurations pointing proximally. The boundary zone of the median layer is commonly marked by a distinct, dark line (PI. 20, fig: le). Endozone — In the recumbent endozone, the zooecial chambers have trianglar cross sections, and become polygonal distally (PI. 20, fig. le). Zooecial growth axes are nearly straight throughout the endozone, diverging from the distal growth axis at a low angle, com- monly less than 30° (PI. 20, fig. 1d). Zooecial walls are thin and parallel-sided with a homogenous to subgranular cortex, and a thin, dark-colored zooecial lining except on the recumbent wall (median layer). Exozone — In the exozone, the large dimorphs are irregular in distribution (PI. 20, figs. la, b). The walls of adjacent large and small dimorphs are subsymmetrically to nonsymmetrically thickened across the boundary zone. The walls of large dimorphs are com- monly nearly parallel-sided. Zooecial chambers have subcircular, elliptical, or subelliptical cross sections. The walls of large dimorphs protrude slightly above the zoarial surface. Small dimorphs are either scattered between large dimorphs in monticular areas, or are concentrated in these areas. Monticular BULLETIN 291 90 apog ‘vadg IZN 66 66 °ZN 66 tI 66 LE 66 ef oT IT T T T N ARO £0° £0° e720" #S ot OT 80° 60° c0° £0° SId}IWIT][IW Up, UGXININ-USSD-YO9Z 6T UQXIN-YS§8D-YOIZ 3) UQXWN-YSSD0-4YO9Z e720" 810° Se UdxW-"4Ss)-YO2Z au0zoxy - sydiow jog []V - [B199007 970° YL-14 Top 10 UL-AZ *X 160 WM-1Z ie IH-1Z [el1e0Z7 LO Iajoe1eyO (SSQHd1OD) FYOdOTYWONPF VIXFLId JO SLNANAYNSVAW AO AUVNWAS TVOILSILVLS el ATaV.L 91 CeRIopoRID CycLosToMEs (Bryozoa): NYE SIOJIWIT| [IU UT, I if Le Le 0 T TGQGIN=7aPZ I Lf Le Le 8£ 610° 0S0° 020° £60° YL-IM9PZPO WOXINN-US80-YO9Z I T Le Le 9 £0" UT OT fT UOXIN-48§)-4O9Z I T Le LZ cl +0° ir 80° a UCXINN-YS80-4O9Z T I LZ Le EE T0° tT or st WdXIN-4S8D-4O9Z auo0zoxy - sydiowAjog adie - [e199007 T T 66 66 0 4 U§899Z/UD-1d PZ Ls I 66 66 St 920° LS0° £10" Lev YL-TM2ZPO (‘uod) su0zoxy - sydiowsjog |[Yy - [8199007 Se aPod ‘vedg IZN 9%ZN N ‘A‘O *S *& «WO Jayoeieyo (SSQUd109) PYOdOTFWONF FIXF.LIG AO SLNAWNAYASVAW AO AUVINWAS IVOLLSLLV.LS BULLETIN 291 92 apog ‘vedg IZN 9ZN Ls LS 'Z-S0tZ INNS ‘Aveo, adAy cy} JeaU WoIZ UsUTIOads palzuap! suo pur :0Z7T AN ad4}0}09TJ ‘T adoo NAWIOddS OL ADH SI9JIUIT [IW UT y UqxXI-4SsD-YD9IZ4IO YL-LM92ZPO UGX N-USS0-4YO9Z UQGXIW-USsSD-YOPZ UGXIWN-YSS0-YO9Z UQGXI-USsSD-YOIZ TsjJoeIeYy) 4 pa aerecce - sydiowAjog [[BWs - [8199007 ft 820° $90° £20° Ltr ST c eT OT 6'T £ 10° 90° 20° 80° $c 20° L0° £0° or auo0zoxg - sydiowAjog [[BUls - [B109007 ‘A'O aS aX «WO (SSAHdIOD) FYOMCOTKWONF FIXF LIC AO SLNANAYXNSVAW AO AUVNWAS TVOLLSILVLS Crerroporip CycLosToMEs (Bryozoa): Nye 93 areas are patchlike in distribution (PI. 20, fig. 1b). The walls of small dimorphs commonly show annular thickenings. The thicken- ings of adjacent small dimorphs are symmetrical across the zooecial boundary zone. Monilar profiles are circular, obovate to inverse pyriform, or less commonly, nonsymmetrically thickened across the boundary zone. The chambers of small dimorphs are commonly elliptical in cross section. Diaphragms — Intermediate diaphragms were sometimes ob- served. The diaphragms are thin (.005 to .008 mm) and planar, and were most commonly observed close to the zooecial aperture, al- though they were sometimes seen as deep as the zone of zooecial flexure. TABLE 14 FREQUENCY OF VISUALLY ESTIMATED OUTLINE OF ZOOECIAL CHAMBERS IN DITAXIA ANOMALOPORA (GOLDFUSS)* Circ. Ellip. Ov. Pyr. Polyg. Triang. Irreg. Exozone - Large Polymorphs Regular 1 7 Sub 10 7 1 Irregular 1 Exozone - Small Polymorphs Regular + 29 5 Sub 8 6 1 Irregular + —_ *Outlines of zooecial chambers were estimated on the lectotype, UB 120 (50 zooecia) and USNM 2405-2 (50 zooecija). Discussion. — The designation of lectotype is consistent with a previous restriction made by Von Hagenow (1851). Goldfuss fig- ured three specimens in his plate 10, figure 5, as Ceriopora anomalo- pora. Von Hagenow considered only the specimen figured in plate 10, figures 5c, d, as D. anomalopora, and refigured this specimen in his plate 4, figure 9c. This restriction was later followed by Gregory (1899, p. 406) and Canu and Bassler (1922, p. 101). The word “Holotypus” is written on a label with the specimen, but the label was most certainly added much later than 1826 by someone other than Goldfuss. Remarks on dimorphism. — Zooecial dimorphism in the exo- zone is expressed in the bimodal frequency distribution of the zooecial 94 BULLETIN 291 751A 75 B ~ ~> oO oO Cc (= ® 250 25 lox Co 2 2 NO on Le) on mm 4d 22. as) 1.0 20 ZcCh-CsSn-NMxDn ZcCh-CsSn-Mx Dn ZcCh-CsSn-NMxDn 754C ~ ‘ 100 (= tb) 5 50 N75 & lox cb) ® > “ PS bYX0) 25 s =] £ 25 pe | oO mm a ve. i 2324 756 CdZcWIl-Th ZdPr-Cn/ZcCsSn Text-figure 10 A-D. Histograms and cumulative curve from the lectotype and one topotype of Ditaxia anomalopora (Goldfuss). A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness. D. Count of interzooidal pores per zooecial cross sectian. CrERIOPORID CycLosToMEs (Bryozoa): NYE 95 diameters (Table 13A), and in the appearance and apparent struc- ture (see below) of the zooecial walls. The general terms, large and small polymorphs, are used for typical members of each dimorph rather than autozooecia, and the like because the characters described are not here considered sufficient to infer function or description of enclosed soft parts. Remarks on microstructure. — Most of the specimens studied, including the lectotype, show poorly preserved microstructure, and in only three specimens were structures preserved well enough to attempt interpretation. The patchy occurrence of nondescript, calcareous material, and the irregular occurrence of a laminar tissue may well be, in part, artifactual and due to diagenetic changes of primary structures. The appearance of the wall in some large poly- morphs is strongly suggestive of lunaria-like deposits. These struc- tures, however, are not here identified as lunaria because of four factors: 1) The structures identified as possible lunarial deposits, although commonly occurring on the proximal side of the zooecium, were sometimes observed in other positions around the zooecial cavi- ties of large polymorphs. Lunarial deposits, typically, are em- placed in the proximal portion of a zooecial wall (Utgaard, 1968a, p- 1033). 2) As seen in tangential section, patches of calcareous tissue, simi- lar in appearance to the possible lunarial structures, were seen in zooecial walls of small polymorphs. 3) The possible lunarial deposits were not associated with any lunaria-like inflections of the outline of the zooecial cavity. 4) In well-preserved fistuliporoids, ceramoporoids, or lichenoporids, the lunaria project orally from the zoarial surface as hoods. These were not seen on any specimens of D. anomalopora. The lunaria-like structure is often missing, or poorly expressed, in shallow tangential section. Sometimes, as seen in longitudinal section, the lunaria-like deposit is capped by indistinct laminate tissue. Possibly the mode of growth changes slightly in the outer exozone, and lunaria-like tissue is not deposited, suggesting onto- genetic control. 96 BULLETIN 291 Genus HAPLOECIA Gregory, 1896 Type species: Millepora straminea Phillips, 1829, by original designation. 1896. Haploecia Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Jurassic Bryozoa, p. US7: 1922. Haploecia Gregory, Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 61, 11953; Hi Gregory, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G71. 1965. Dendroecia Cotillon, and Walter, Soc. Géol. France, Bull., vol. 7, ser. 7, pp. 934-935. Tentative diagnosis.—Zoaria_ branched; distally growing branches intersect and anastomose. Branches have coaxial endozones and exozones. Zooecia intersect zoarial surface obliquely. Branches commonly encrusted by one or more intrazoarial overgrowths. In endozone, zooecia with smaller diameters often surround single large zooecium growing parallel to distal growth axis of branch and located close to center of branch. In exozone, cortex composed of light-colored, indistinctly laminated, calcareous tissue. Laminae slightly crenulate and, in general, broadly arched, convex orally. Lamination commonly be- comes obscure or disappears in outermost cortex. Zooecial linings dark-colored, with crenulate, longitudinally directed laminae. Lateral spinelike extension of light-colored cortex tissue sometimes extends into zooecial chamber as mural spines, or sometimes submerged beneath thick deposits of zooecial lining. All zooecia distal to unencrusted growing tips sealed by simple external walls. Intermediate and basal diaphragms occur but not common. Taxa included. — Based on the examination of primary types, two species are here considered to be correctly assigned to Haploecia: Ceriocava multilamellosa Canu and Bassler, 1922 [Lower Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland], and the type species, Millepora straminea Phillips, 1829 (Middle Jurassic, Yorkshire, Eng- land). Internal characters of other species assigned to Haploecia are unknown to me. Discussion. — On the basis of wall structure and occurrence of diaphragms, Haploecia is retained in the cerioporids. The occurrence CrERIOPoRID CycLosTomeEs (Bryozoa): NYE 97 of simple external walls and attendant modification of growth habit is known to occur in only one other cerioporid genus, Diplocava. Simple external walls resemble externally the porous, foraminate, apertural seals seen in the Salpingina. Viskova has recently studied genera assignable to the Salpingina and has redefined three families on the basis of internal characters (Viskova, 1965, 1968). Assign- ment of Haploecia would be consistent with the revised concept of the Eleidae d’Orbigny except for differences in wall structure. Vis- kova (1968, p. 175) reported that zooecial walls in eleids are longi- tudinally fibrous. In specimens of Haploecia available for study, wall structure was poorly preserved. Orally convex lineations (PI. 23, fig. Ic; Pl. 26, figs. 1b, c; Pl. 28, fig. 3b), however, are interpreted as remnants of original lamination. Gregory (1896, p. 157) allied Haploecia with the entalophorids, and in 1899 (pp. 288-9) suggested that Haploecia was an early stage in the evolution of the Eleidae from the Entalophoridae. The meli- cerititids, a later evolutionary stage in Gregory’s scheme, are charac- terized by tubular zooecia, but are reputed to possess both opercular and avicularian structures (Gregory, 1899, pp. 287-292; Levinsen, 1912, pp. 10-13, 19; Bassler, 1953, p. G75). The occurrence of struc- tures presumably diagnostic for cheilostomes (Bassler, 1953, p. G147) in tubular bryozoans led Gregory (1899, p. 287) to state that “The Cretaceous family, the Eleidae, is important, as it breaks down the distinction between the Cheilostomata and Cyclostomata on these characters”. The external similarity of the type species of Haploecia to some Cheilostomata is striking (Pl. 23, fig. 1b). As observed by Gregory (1899, p. 288), “... the zooecia, at their distal ends, are hexagonal, bounded by ridges, and have a small subterminal aperture at the upper part. This arrangement is very similar to that of the genus Cellaria.” This similarity, however, is due to convergence, as can be ascertained by reference to internal structures. The apertural structure resembling a porous frontal wall is a simple external wall. The zooecia in Haploecia are tubular, rather than boxlike; and the zooecial walls in Haploecia are compound, being shared with ad- jacent zooecia rather than the cuticle-lined individual cases typical of the Cheilostomata. 98 BULLETIN 291 Canu and Bassler (1922, pp. 97-8) were apparently the first to study thin-sections of specimens assignable to the type species. They observed the enlarged axial zooecium, the simple external walls (“facettes”), the absence of mesopores, and the general ab- sence of other diaphragms. They referred Haploecia to the Cerio- cavidae. As presently understood, Certocava and Haploecia exhibit large morphologic differences, although the mode of growth is similar. Both have coaxially arranged endozones and exozones, and show increased diameters of zooecial chambers from endozone to exozone. In Certocava, however, zooecia apparently continued to grow in the exozone for much of the life of the colony; in Hafloecia, orally di- rected growth terminates relatively soon after zooecia become exo- zonal in character. Terminal diaphragms are emplaced in Ceriocava, simple external walls in Haploecia. Basal diaphragms are numerous and closely spaced in Ceriocava but were rarely emplaced in Haploecia. Anastomosis occurs in Ceriocava but is associated with the bifurcation of two branches rather than with the intersection of their growing tips as seen in Haploecia. Cotillon and Walter (1965, pp. 934-5) erected Dendroecia be- cause of the nature of the budding and the appearance of the ovicells in specimens they assigned to Ceriocava multilamellosa (Canu and Bassler, 1922). The internal morphology of the primary types of Ceriocava multilamellosa is consistent with the generic concepts of Haploecia, and Dendroecia is determined to be a junior subjective synonym. Haploecia straminea (Phillips), 1829 Pl. 23, figs. la-f; Pl. 24, figs. 1a-f; Pl. 25, figs. la-d, 2a-b; Pl. 26, figs. la-d 1829. Millepora straminea Phillips, Illustrations of the Geology of Yorkshire. Description of the strata and organic remains of the Yorkshire Coast, pp. 144, 149, pl. ix, fig. 1. 1893. Pustulopora straminea (Phillips), Gregory, Yorkshire Philos. Soc., Ann. Rept., p. 60, text-fig. 2. 1896. Haploecia straminea (Phillips), Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Jurassic Bryozoa, pp. 159-161, text-fig. 12, p. 160. 1922. Haploecia straminea (Phillips), Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 61, pp. 97-8, pl. 14, figs. 14, 15, text-fig. 25, p. 98. 1945. Haploecia straminea (Phillips), Melmore, Catalogue of Types and Figured Specimens in Geological Department of Yorkshire Museum, p. 216. Type.— The lectotype is YM-T81/2 (figured by Gregory, CERIOPORID CycLostoMEs (Bryozoa): NYE Se 1893, text-fig. 2, p. 60, and 1896, text-fig. 12, p. 160) by indication, Gregory (1896, explanation of text-fig. 12, p. 160). Type locality and horizon. — The label with the type specimens bears only information bearing on Gregory’s publication, and no locality data are given. Phillips (1829, pp. 144, 149) cited the localities as “Scarborough (very rare), from the Cornbrash; Gris- thorpe, Cloughton, Owlton, Crambe, Westow, Ellerker, from Gray Limestone or Bath oolite, England.” Sediments bearing fragments of bryozoans identified with H. straminea exposed at these localities are questionably referred to the Bajocien (pers. comm. J. W. Neale). Material studied. — The lectotype YM-T81/2 and paralectotype YM-T81/1 were borrowed from the Yorkshire Museum. Thin-sec- tions and acetate peels were made from both the lectotype and paralectotype. In addition, an encrusted specimen of H. straminea was re- vealed in a thin section of Ceriocava corymbosa (Lamouroux) [USNM 32181-2, from the Bathonien, Langrum (Calvados), France]. Acetate peel replicas of all specimens sectioned are pre- served in the collections of the National Musum of Natural History and of the author. Description. — Growth habit —Zoaria are profusely branched with branches growing distally in all directions (PI. 23, fig. la). Branches inter- sect and anastomose in the proximal portion of the colony forming a densely matted network. Branches have subcircular to elliptical cross sections and wedge-shaped to rounded growing tips. Endozone — Zooecial walls are thin and slightly undulatory (PI. 24, fig. 1c). Generally, the walls are nearly parallel-sided, but prominent asymmetrical thickenings are sometimes observed. The walls are light-colored and granular. Zooecial linings are thin or missing in the inner endozone (PI. 23, figs. 1c, d, f; Pl. 24, fig. Ic). Interzooidal pores are rarely seen. Exozone —Near the tips of unencrusted primary branches, zooecial walls are sometimes thickened asymmetrically across the zooecial boundary zone (PI. 23, fig. 1d). Walls have submoniliform profiles because of moderate variation in thickness longitudinally (Pl. 23, fig. 1d) and because of the relatively wide flare of inter- zooidal pores (PI. 23, figs. 1c, d, e). The outer exozone portions of 100 BULLETIN 291 zooecial walls are thicker in a gradient directed proximally from un- encrusted primary branch tips (Table 15) in at least some encrusted primary branch tips (PI. 24, fig. la), and in intrazoarial over- growths (PI. 24, fig. 1b). Thicker zooecial walls generally are nearly parallel-sided because of in-filling of thinner portions of the wall, and because of reduction in the “flare” of interzooidal pores (PI. 26, figs. Ic, d). Zooecial cross sections are generally subelliptical (Table 16) except at apertural level. At the apertural level, the wall thins rapidly to a rimlike extension (PI. 23, fig. 1c; Pl. 25, fig. 2b; Pl. 26, fig. 1b), and zooecia have polygonal, commonly hexagonal outlines (Pl. 23, fig. 1b; Gregory, 1896, text-fig. 2, p. 60). Simple External Walls — External walls were observed in nearly all zooecia more than about 1 mm from unencrusted branch tips. The exterior walls are thicker and generally appear more robust in a gradient directed proximally from unencrusted branch tips (com- pare PI. 23; figs!le, d, and Pl°25) fies! 2a, b, to Pl. 24) figs: lev rand PI. 26, figs. la-d). Peristomes are low near unencrusted branch tips (Pl. 23, fig. 1b); but proximally from branch tips, peristomes may extend as much as .2 mm above the zoarial surface (PI. 24, fig. 1f; Pl. 26, figs. 1b, c, d). Peristomes have circular to elliptical cross sections and are located in the centro-distal portion of each zooecium (Pl. 24, figs. le, f). Measurements of the maximum dimensions of peristomial cross section (the restricted skeletal aperture) and cross sections of diaphragmal pores are summarized in Table 15. Diaphragms — Both intermediate and basal diaphragms (PI. 25, fig. 1b) occur but, in general, are not numerous. Diaphragms em- placed in the almost wholly recumbent zooecia of some interzoarial overgrowths are attached, in part, to simple external walls (Pl. 26, fig. 1b). In addition, basal diaphragms were sometimes observed in peristomes (PI. 26, fig. 1b). Discussion. — The original figure by Phillips (1829, p. 19) is generalized and of little use in identifying other specimens with H. straminea. As noted by Gregory (1896, p. 160). “This species was figured so imperfectly by Phillips that it had been variously in- terpreted by foreign authors”. Gregory refigured specimen T81/2 twice (1893, text-fig. 2, p. 60; 1896, text-fig. 12, p. 160); both figures are somewhat idealized illustrations showing regularly arranged hexa- 101 CrerioporIp CycLostomeEs (Bryozoa): NYE ‘sdyy youviq pojsn:ouaun woz WW ¢ URY} ssaTt SIOJIUNT| [IW UTy I if LT LI 0 ‘Y USsD9Z /YO-1d PZ I if LI LI Ot c0" $0° z0° 80° YL-TM9IZPO T I LI LM Le v oT TT 6c WQXWN-YSSD-YO9Z UQGXIN-US8D-YO9Z I I LI HN 4 +0° Sk 60° I UWQXINN-YS8)-YO9Z T T LI js) st £0° co [A 8c UQXIN-USsSD-Y4D9Z T gUOZOXY - [8199007 c c “82 ‘09 WM-IZ [ele0Z7 2poy 99dg) IZN (OZN N AND) 4S 4X * WO TSE SE US) (SdITIIHd) VFANINFALS FIDAOTIVH AO SLNANAUNSVAW JO AUVWWAS TIVOLLSLLV.LS st ATHVL BULLETIN 291 102 ‘sdi} youvig pajsniduaun wolf Wu ¢ UBY} a1oW 10 ‘sd1} yOURIG paysni0Ug * ‘sd youesg pajsniousun wor Wu ¢ UBY) SsaTt SId}JIUWIT[[IW UT, I T 8 3 $2 Z0" or 90° et WL-LMPZPO UGsINN=US§OatOeZ i if 8 8 IT c LT sT Ve UQXI-US§)-YOIPZ I I 8 8 $I c0° or 80° vr UCXWN-US89-YD9Z I I 8 8 eT ZO" 8st 9 0c UQXIN-US8)-YOIZ 8 QUOZOXY - [8199007 T I oT 91 Le Z00° $00° £00° 800° IqQuN-1dPZ T I Lt LI 9b 200° £00° 000° $00" HL-FIIM9Z (‘uod) t auozoxg - [B199007 apop 9adg_ — IZN (OZN N ‘ACO «S aX «WO TSE TE) (SdITIIHd) PFANINFULS FIDAOTAIFH JO SLNANAANASVAW JO AUVNWWAS TVOILSILVLS 103 Cerroporip CycLosToMEs (Bryozoa): Nye ‘T/I8.L WA :adAjoyoa[eieg ‘¢ ‘T/I8.L WA :2dAjoyajered pue addjoja7T “Zz ‘7/I8.L WA 2 9dAjo}27T “T adOoO NAWIOddS OL AIM ‘sdij youersq pajsniouaun WolJ Wu ¢ UY} a10UI JO ‘sd1} YOURIG pajsnioUg * SIOJIWT|[IW UT, 4 Z L 100° 900° YL-PISa suseiydeiq I I oT oT LE 200° $00° £0C° 800° UqxW-USs-dpsd-M aS 4 c £1 at 9€ cc0" £90 80° LL0° UCXIN-USsO-MAS HeAA [Butaxq afduig Lt T Of Of 0 ‘T U$899Z/UD-1dPZ I T 0€ Of 6£ $00° TO T0° £0° UL-LM9ZPO UGXWN-4S80-YO9Z T T Of Of 1&4 o oT OT v~ UGXIN-YS8D-YDIZ I T Of Of 6£ 20° L0° TO cr UQXINN-YS80-YO9Z I T 0€ Of 6£ +0° or ZO" Lv UQXIN-USsD-YOIZ auozopuy - [8199007 I T 8 8 ce L00° $c0° c10° $£0° GL-F TMZ I I 8 8 0 4 US899Z/4D-1d PZ (‘u0d) *9U0zZ0xq - [8199007 apod ‘vedg TZN 9ZN N “AO «S «aX «XO TOIELES (SdITIIHd) FANINVULS FIDAOTAFH AO SLNAWAYASVAW AO AUVWWAS IVOLLSLLV.LS 104 BULLETIN 291 75,4 75 B > ~ oO oO (= = o 250 250 o rou C : 25 25 mm 4a 2 a 1.0 20 ZcCh-CsSn-NMxDn ZcCh-Cs Sn-Mx Dn ZcCh-CsSn-NMxDn 754C = 100 oO (= o 5 50 N75 D rox o = > _ = 450 25 M4 = e 25 S oO mm | a2 1; 2°33: 4 5-6 CdZcWI!-Th ZdPr-Cn/ZcCsSn Text-figure 11 A-D. Histograms and cumulative curve from the lectotype and paralectotype of Haploecia straminea (Phillips). A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness. D. Count of interzooidal pores zooecial cross section. Cer1oporip CycLostomes (Bryozoa): Nye 105 aS frequency NO .02 .04 .06 thickness in mm. Text-figure 12 A-C. Histograms showing variation in thickness of peri- stomial diaphragms in Haploecia straminea (Phillips). A. More than 3.0 mm from tip of encrusted branch in YM T81/1. B. More than 1.2 mm, less than 3.0 mm from branch tip in YM 1T81/2. C. Less than 1.2 mm from tip of un- encrusted branch in YM T81/2. 106 BULLETIN 291 TABLE 16 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN HAPLOECIA STRAMINEA (PHILLIPS) Cire, Elilip: Ov. Pyr. Polyg. Triang. Irreg. Exozone Regular 1 1 1 Sub 11 2 Irregular 1 Endozone Regular 1 Sub 11 4 1 + Irregular 3 1 1 gonal zooecial apertures sealed by simple external walls. In the ex- planation, Gregory said that the figure is, “Part of the type speci- men of Haploecia straminea (Phil.)”. Haploecia multilamellosa (Canu and Bassler), 1926 Pl. 27, figs. 1a-e, 2a, b, 3a-d; Pl. 28, figs. la-e, 2, 3a, b; Pl. 29, figs. la-e, 2a-c, 3; Pl. 30; figs: 1a,.b; 2, 3;-Pl.31 Migs: las byZaep 1926. Ceriocava multilamellosa Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 67, pp. 68-70, text-fig. 35A-F, p. 69, pl. 9, figs. 1-10. 1965. Dendroecia multilamellosa (Canu and Bassler), Cotillon, and Walter, Soc. Géol. France, Bull., ser. 7, vol. 7, pp. 934-5. Type. — USNM 69922-1 is designated as the lectotype. This specimen was figured by Canu and Bassler (1926, pl. 9, figs. 1 — middle specimen, 5, 6), and here (PI. 27, figs. la-e). Type locality and horizon. — Lower Cretaceous (Valangian): Sainte Croix (Vaud), Switzerland. Material studied. — Only the lectotype and single paralectotype of the five specimens originally figured by Canu and Bassler (1926, pl. 9, figs. 1-10) have been found in the National Museum of Natural History collection. Both specimens bear the label “cotype.” The lectotype, USNM 69922-1, is the specimen figured by Canu and Bassler (pl. 9, figs. 1 — the middle specimen, 5, 6). Three thin- sections and four peels on one acetate slide were prepared from the lectotype. Paralectotype specimen USNM 69922-2 is the specimen figured in Canu and Bassler (1926, pl. 9, figs. 1 — second specimen from the right, 7 and 8). Four unnumbered thin-sections, labeled type and prepared by R. S. Bassler, are considered paralectotypes. CERIOPORID CycLostomEs (Bryozoa): NYE 107 The thin-sections were figured by Canu and Bassler (text-fig. 35, p- 69) as follows: No. 41, text-fig. 35B, C; No. 41.5, text-fig. 35D; No. 41.7, text-fig. 35E; No. 41.6, text-fig. 35F. Thin-sections and acetate peels were made from nine topotypes identified by R. S. Bassler. These bear the number USNM Loc. 2384. Description. — Growth habit — Branches are roughly cylindrical, consisting of a primary branch with a well-defined coaxial endozone and up to three thin (Table 17, Ov-Th) intrazoarial overgrowths. Zooecial apertures are generally arrayed in parallel ranges. Zooecia in ad- jacent ranges commonly alternate in position longitudinally (PI. 27, figsslby 3b, d> PI 28: fie. 3a; Plo 29; tig. 1b): Distally growing branches sometimes intersect and anastomose (Pl. 29, figs. la, 2a). Where the plane of intersection is nearly normal to the direction of zooecial growth, each zooecium is generally sealed by a thin, nonporous diaphragm (PI. 29, figs. 1c, d, 2b, lower center, 2c). Where the plane of intersection is oblique to zooecial growth, the zooecial walls of intersecting zooecia merge and continue to grow orally (PI. 29, figs. 2a, b, upper center). Endozone — Zooecial walls are parallel-sided and occasionally have rounded to spinose projections (PI. 28, fig. 1d). Longitudinally, the walls are moderately undulatory (PI. 28, figs. la, d; Pl. 29, fig. Ic); Exozone — Zooecial walls have submoniliform profiles and are symmetrical across the zooecial boundary zone. Zooecial cross sections are commonly elliptical. In the outer exozone, mural spines are sometimes numerous within individual zooecia. The spines have light-colored (subgranular) cores which contrast with the dark- colored laminated tissue of the zooecial lining (PI. 28, figs. lc, d; Pl. 31, fig. 2b). Interzooidal pores are nearly cylindrical and only slightly flared. Simple External Walls — External walls were observed in all zooecia subjacent to intrazoarial overgrowths. The walls are rela- tively thin and composed of dark-colored, wavy, laminate tissue similar in appearance to the zooecial lining (Pl. 30, figs. la, b). Laminae were sometimes observed to flex aborally and merge with the zooecial lining (PI. 30, figs. 1a, b; Pl. 31, figs. la, b, 2b). Peri- stomes are commonly subcircular, located centrally, and extend only BuLLETIN 291 108 SIIJIWN [IW Uy z £ I+ It 0 8 9Z /UD-dgoZ 6 zZ +E +£ I+ 500° £10° £00° 920° YL-UTIMIZ Z € I+ It 0 9 USsD2Z/UD-1d PZ + ST ST 9€ £0° 60° $0" 91° AIT -YL-1M9ZPO £ 9 91 LE LO" 81° 80° 92° S3UT-Y,L-[M9ZPO UGX N-USSD-YD9Z Z € It It 61 + 0% +1 £'€ UQXIN-USsD-YD2Z Z £ I+ It 8I Z0° 80° S0° jae UGX N-4S8D-YD2Z Z £ Iv It ST z0" St or’ 02° UGXIN-USSD-YDIZ d9uUOZOX - [B199007 T L +I $$ 60° AV er 97° UL-A0 I L L cz bas 01 3; i UQXIN-USsD-1G1d I L L +2 ¢" Lit ct +2 UqxXW-USsD-1g [BIIB0OZ apop9edg_ = IZN-OZN N UN) #8 #X *WO Joyoereyy (YHISSVE GNV ANVO) FSOTTAWFTILIAW FIDTOTIFH AO SLNAWANASVAW AO AUYVWANDAS TVOILSILLV.LS Lt ATa#VL 109 NYE CERIOPORID CycLosToMEs ( Bryozoa) (OL) b-P8EZ “(ST) I-t8EZ 907 IWNSA :49\sseg Aq palznuapt saddjodoy, (6) 1-27669 WNSQ 244101007 (02) c-+8E2 INSN ‘4a[sseg Aq patznuap! edsjodoy, - (+) G-P8EZ “(+) T-+8E2 WNSA :Je[sseg Aq parynuap! sadAjodo 7, (9) b-P8EZ “(ST) C-F8EZ “(41) I-b8EZ 907 WWNSA :4a[sseg Aq sadAjodoy, § (2) 1-27669 WNSA 244103007 "(€) [-t8E% “907 WNSN ‘49]sseg Aq parznuapr adAjodoy, “(ZL) I-+8€% “907 WNSA :42[sseg ‘S ‘y Aq palzuuap! uauroads addjyodoy, £(¢) 1-27669 INNSD ad 4}099aT “(41) T8207 IWNSA :2I8Seg ‘S “Y Aq palzuap! suaunsads adAyodoy, $(Z) 1-22669 WNSA 2dAr01007 (ZI) b-48EC (ST) I-+8E% “207 WNSD :4a[sseg “Ss ‘y Aq parynuapr saddjodoy, § (+1) 1-22669 NNSA adAz0309T Nn tH ON OH "(yoea T) 6-+8EZ ‘B-P8EZ “L-b8E7 “b-b8EZ ‘Z-+8EZ ‘I-b8EZ “90T WNSD ‘4alsseg “§ ‘y Aq parynuap! sadfjodoy, ! (1) 1-27669 WNSQ 244103027 “1 dd0O0 NYWIOddS OL AIM SIDJIUIT[[IW UT, 8 I + 02 SZ Z00° 800° $00" E10" UqxW-USsD-dpsq-MaS L Z 8 8 LI 10" 120° SSO" 780° UQXIN-USSD-3Sd-M AS 9 + eb cb tt 900° +10" 900° LZ'0 4YL-MdS SI[EAA [BUtaxq arduis ¢ I 3 g 800° 910° IGuW-1dPZ I dL iL L LZ £0° 1 80° 91° UQXIN-USsD-YO2ZUD auozopuy - [8109007 S I 3 € £00° 010° IGU-1dPZ apo) 22d§ = IZN OOZN N xD *S #X TopeEIEYD (YHISSVA UNV ANVO) FSOTTANFTILIAW FIDZOTIFH JO SLNANAUOSVAW JO AUVWWAS IVOLLSILLVLS 110 BULLETIN 291 slightly above the zoarial surface. Peristomial apertures subjacent to overgrowths are commonly sealed by dark-colored laminate tissue continuous with the diaphragm (PI. 31, figs. Ib, 2a): Diaphragms — Intermediate diaphragms (Pl. 30, fig. 3) were observed rarely in primary branches. The diaphragms are planar to strongly convex orally and merge continuously with the zooecial lining. Diaphragms are moderately thin (Table 17, IntD-Th). Intrazoarial overgrowths — Zooecia bud obliquely to the basal layer and intersect the zoarial surface obliquely; commonly, little or no zooecial bend is seen (PI. 28, fig. 1a; Pl. 29, figs. la, c). Al- though zooecia are relatively short, both thin-walled (endozonal) and thick-walled (exozonal) portions are present. Mural spines sometimes project from the basal layer (PI. 30, fig. 1b; Pl. 31, fig. 2b). Intermediate diaphragms were rare to moderately numerous. At the most, each zooecium contained a single diaphragm. Discussion. —Canu and Bassler, in their original description of Ceriocava multilamellosa, noted that “The appearance of facettes in the genus Ceriocava still remains a mystery.” Cotillon and Walter (1965, p. 935) recognized other morphologic characteristics that were not compatible with their concept of Ceriocava based on the type species, C. corymbosa (Lamouroux). Among these differential characteristics in C. multilamellosa were the lack of transverse diaphragms and mesopores, zooecia less bent, increase in zoarial diameter by overgrowth, and very different appearance of the ovicell. TABLE 18 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN HAPLOECIA MULTILAMELLOSA (CANU AND BASSLER) Circ. Ellip. Ov. Pyr. Polyg. Triang. Irreg. Exozone Regular 31 7 Sub 2 lrregular 1 Endozone Regular 1 12 4 Sub 7 14 4 3 Irregular 1 Cerroporip CycLostomes (Bryozoa): NYE 111 7TH A 75 B ~ ~ oO oO [= ts @ ®50 25 oa oOo 2 ® 25 25 mmi 2. .3 1.0 20 ZcCh-CsSn-NMxDn ZcCh-CsSn-MxDn ZcCh-CsSn-NMxDn Cc >25 5 100 Cc o 3 ~ 75 fo 5 @ na > ok = 50 25 = 3 £ 25 a3 oO mm A 72 162.34 5).6 CdZcWI-Th ZdPr-Cn/ZcCsSn Text-figure 13 A-E. Histograms and cumulative curve from the lectotype and two topotypes of Haploecia multilamellosa (Canu and Bassler). A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness as measured in longitudinal section. D. Compound zooecial wall thickness as measured in transverse section. E. Count of inter- zooidal pores per zooecial cross section. D2 BULLETIN 291 Cotillon and Walter described the appearance of ovicells as seen in specimens collected from the calcaire lumachelliques (Hauterivien and Barremien) from localities in the Basses-Alpes, France, which they assigned to C. multilamellosa. On the basis of these differences in morphology, Cotillon and Walter erected the new genus Den- droecia for C. multilamellosa Canu and Bassler. Based on an examination of thin-sections of syntypes and topo- types identified by Canu and Bassler, C. multilamellosa has charac- ters consistent with the tentative diagnosis of Haploecia and is assigned to Haploecia. H. multilamellosa differs from the type species, H. straminea, in having branches with smaller diameters and in the more regular arrangement of zooecial apertures. Zooecia have smaller cross sec- tions and fewer interzooidal pores per zooecial cross section, but more mural spines. Peristomes commonly are located centrally with relation to the zooecial walls. In H. multilamellosa, zooecia in over- growths generally grew obliquely away from the basal layer; in the type species, zooecia usually grew recumbent to the basal layer for much of their length. Genus HETEROPORA Blainville, 1830 Type species: Ceriopora cryptopora Goldfuss, 1826, by subse- quent designation, Gregory (1896, p. 201). 1830. Heteropora Blainville, Zoophytes: Dictionnaire de Science Naturelles, vol. 60, p. 381. 1834. Heteropora Blainville, Manuel d’Actinologie ou de Zoophytologie, p. 417. 1851. Pars Ceriopora Goldfuss, Von Hagenow, Die Bryozoen der Maastrichter Kreidebildung, p. 53. 1896. Heteropora Blainville, Gregory, Catalogue of Fossil Bryozoa in the De- partment of Geology, British Museum (Natural History), The Jurassic Bryozoa, p. 201. 1909. Heteropora Blainville, Gregory, Catalogue of Fossil] Bryozoa in the De- partment of Geology, British Museum (Natural History), The Cretaceous Bryozoa, vol. 2, p. 185. 1933. Non Heteropora Blainville, Borg, Zool. Bidrag fran Uppsala, Band 14, Ppiezoos 1953. Heteropora Blainville, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G66. Tentative diagnosis. — Zoaria massive to branching, composed of superposed intrazoarial overgrowths. Endozones thin, directly adjacent to basal layer, or not developed. Zooecial walls granular, becoming indistinctly laminate in outer CerRIoporIp CycLosToMEs (Bryozoa): NYE HS exozone. Laminae broadly arched, convex orally and continuous across zooecial boundary zones. Thin zooecial lining commonly pre- sent in endozone and exozone. Intermediate diaphragms common, basal diaphragms occurring rarely. Taxa included. — Only the type species, Ceriopora cryptopora, is here included. Heteropora dichotoma, included in Heteropora by Blainville, is here considered to be correctly assigned to Ditaxta. Characters seen in thin sections of specimens assigned to H. dicho- toma by Bassler, the third species included in Heteropora by De Blainville, are not consistent with the concept of Heteropora as understood here. Thin sections made from topotypes of H. magna O’Donoghue and O’Donoghue and H. pacifica Borg, and sections made from a syntype and identified specimens of H. pelliculata Waters, revealed the following characters: 1) Zoaria have coaxial exozones and endozones. 2) Zooecia are dimorphic. 3) Zooecial walls are composed of aborally oblique laminae. 4) Terminal (and possibly peristomial), intermediate, and basal diaphragms occur. These characters are not consistent with the concept of Heteropora, and the above species are assignable to another, probably unnamed genus. The internal characters of numerous other species assigned to Heteropora by authors are presently unknown. Discussion. — Heteropora was erected by Blainville for three species first described by Goldfuss (1826) and placed in his genus Ceriopora. Goldfuss (1826, p. 32) had erected the genus Certopora and assigned 28 species to it, all of them new. Heteropora was erected by Blainville to separate from Ceriopora Goldfuss those species which “se distingue essentiellement par l’extence de deux sortes de cellules ou de pores, les unes deux ou trois fois plus grandes que les autres . . . et composées de couches enveloppants”. Blain- ville, in erecting Heteropora, apparently had based his diagnosis only on the figures and descriptions of Goldfuss, and had not ob- served a specimen assignable to the genus because he wrote, “Nous ne voudrions cependent pas assurer ce dernier point, n’ayant pas 114 BULLETIN 291 encore analysé nous-méme une espéce d’heteropore” (“ce dernier point” refers to “de couches enveloppants” Blainville, 1834, p. 417). Gregory was the first reviser of Heteropora Blainville because in 1896 (p. 201) he cited H. cryptopora (Goldfuss) as the type species of Heteropora. In 1909 (p. 185), he noted that “Of these three species H. cryptopora was mentioned first and of the others the C. anomalopora, Goldf., is a Ditaxia and C. dichotoma, Goldf., is a Sparsicavea.” A single character, zooecial dimorphism, had been consistently cited by authors, except Haime (1854), as diagnostic for Heteropora. The occurrence of dimorphic zooecia was not, however, confirmed in this study. “Polymorphism is discontinuous variation in_ the morphology of zooids arising at the same astogenetic stage”. ( Board- man, Cheetham, and Cook, 1970, p. 9). The authors in the discussion following described ways in which dimorphism may be expressed, and these are summarized below: 1) The possession or lack of a given structure. 2) Particular location within a given budding pattern. 3) Difference in size. 4) More complex differences in terms of both structure and func- tion. Consistent differences between zooecia were not recognized in external or internal examination of type and nontype specimens of H. cryptopora, the type species. There was some variation in the size of zooecial apertures; but no consistent arrangement of larger or smaller polymorphs in the sense of Borg was recognized. A diagram of the frequency distribution of zooecial void diameters was con- structed from measurements of randomly selected zooecia in the lectotype and in a single nontype specimen (Text fig. 14A). The diagram shows a continuous and nearly normal distribution with moderately negative skewness. Gregory (1896, p. 201; 1909, p. 185) referred to the small dimorphs as mesopores and stated that Heteropora was “most closely allied to the genus Heterotrypa”, a Paleozoic trepostome. In- ternal characters as outlined here, however, make as close a relation- ship to the Trepostomata as suggested by Gregory appear highly improbable. Borg (1933, p. 283) apparently based his diagnosis of CERIOPORID CycLosToMEs (Bryozoa): NYE 115 Heteropora on Recent species which he assigned to the genus, in part characterizing the genus as “autozoids . . . not forming clusters; kenozoids smaller and much more numerous than the autozoids, lo- cated between them and thus separating them.” The consistent citation of dimorphism as a diagnostic character apparently stems, in most instances, from three factors: 1) Reference to the original figures of Goldfuss which were some- what idealized, especially magnified views. 2) Reference to Blainville’s definition which, again, was derived by examination of figures rather than specimens. 3) Consideration of the characters of species other than the type species. Remarks on wall structure. — The microstructure in all speci- mens sectioned was poorly preserved. In thin section, the zooecial walls are light grey to light brown in color, and homogeneous to subgranular with a scattering of small dark grains. Often the boundary between the zooecial wall and calcite infillings of the zooecial chambers was poorly defined (PI. 33, fig. 2a). Orally con- vex lineations were, however, sometimes observed (PI. 35, figs. 1a, 2a). These are interpreted as remnants of originally laminated microstructure. Light-colored, homogeneous calcite forms irregular bodies in the cortex. This material is probably secondary in origin because it cuts across laminate microstructure unconformably (PI. 35, fig. 2a). The original zooecial wall tissue is inferred to have been laminate. Heteropora cryptopora (Goldfuss), 1826 Pl. 32, figs. la-f; Pl. 33, figs. 1, 2a-c; Pl. 34, figs. 1a, b, 2a, b; Pl. 35, figs. 1a-c, 2a, b 1826. Ceriopora cryptopora Goldfuss, Petrefacta Germaniae, vol. 1, p. 33, pl. 10, figs. 3a-d. 1830. Heteropora cryptopora (Goldfuss), Blainville, Zoophytes: Dictionnaire de Science Naturelles, vol. 60, p. 382. 1834. Hetcropora cryptopora (Goldfuss), Blainville, Manuel d’Actinologie ou de Zoophytologie, p. 417, pl. 70, fig. 4. 1846. Heteropora cryptopora (Goldfuss), Michelin, Iconographie Zoophytolo- gique, description par localites et Terrains des Polypiers Fossiles de France et pays environnants, p. 3. 1851. Ceriopora cryptopora Goldfuss, Von Hagenow, Die Bryozoen der Maastrichter Kreidebildung, p. 53, pl. 5, fig. 6. 1933. Non Heteropora cryptopora (Goldfuss), Borg, Zool. Bidrag fran Up- psala, vol. 14, p. 283. 1953. Hetcropora cryptopora (Goldfuss), Voigt, Geol. Staatsinst. Hamburg, Mitt., vol. 22, pp. 62-3, text fig. 1, pl. 6, fig. 5. 116 BULLETIN 291 Type. — UB 118a is here designated as the lectotype. UB 118a was figured by Goldfuss, 1826, pl. 10, fig. 3a; Von Hagenow, 1851, pl. 5, fig. 6; Canu and Bassler, 1920, text fig. 222A, p. 681; and here, Pli325 figs. a-f. Pi. sayig. Ae Type localty and horizon. — Goldfuss (1826, p. 33) cited the locality as “Petersberge be: Maastricht”. Rocks exposed there are Upper Cretaceous, Maastrichtian in age. Material studied.— Five syntypes were borrowed from the Institut fiir Palaeontologie, Universitat Bonn, Bonn. Six thin-sec- tions and six acetate peel replicas on two slides were made from the lectotype, UB 118a. Three thin-sections and three acetate peel replicas were made from one paralectotype numbered collectively with four other paralectotypes as UB 118b. The paralectotype sec- tioned was figured by Goldfuss, 1826, pl. 10, fig. 3b; and here, PI. 33, figs. 2a-c, Pl. 34, fig. 1b. Most of both specimens remain as remnants after thin-sectioning. Thin-sections and acetate peel replicas were also made from 11 specimens collected from the Maastrichtian, Maastricht, Nether- lands (USNM Loc. 2387). Description. — Growth habit — Zoaria are commonly robust masses with bul- bous outgrowths or subcylindrical branches (PI. 32, fig. la). Over- growth units are lenticular and commonly have moderately to strongly convex distal and concave proximal surfaces (PI. 32, fig. fs Pli33:cties 2b. PL 34 otigs12b). Branches aresfomned Re the Matas expansion of intrazoarial overgrowths from independent loci (Pl. 33, fig. 2b). The chambers of a few zooecia (commonly located near major growth axis) are continuous from subjacent to suprajacent overgrowth (PI. 33, figs. 2b, c; Pl. 34, fig. 2b), in- dicating a boundary between growth phases. The zooecial walls of continuously growing zooecia commonly show thin dark zones fol- lowed by thin-walled and often offset growth (PI. 34, fig. 2b). Laterally, the subjacent zoarial surface is draped by a thin (about .005 mm), dark-colored basal layer which generally sags partly into the subjacent zooecial chambers (PI. 34, fig. 1a). The basal layer is distinctly laminate with laminae directed about parallel to the sur- face of the basal layer. CERIOPORID CycLosTomEs (Bryozoa): NYE itil 7/ Endozone — Zooecia have thin (about .01 mm), parallel-sided walls (Pl. 34, fig. la). The walls are homogeneous to subgranular with thin zooecial linings. Interzooidal pores were rarely observed. Exozone — Zooecia are nearly straight, gently curved, or slightly undulatory in growth. Zooecial walls are symmetrical in thickness across the zooecial boundary zone. The walls are nearly parallel- sided, and show a slight but regular increase in thickness orally. The walls thin near the aperture, and commonly have rounded to acutely lanceolate profiles (PI. 33, fig. 2a; Pl. 34, fig. la; Pl. 35, figs. 1b, 2a). Less commonly, slight variation in wall thickness oc- curs producing submoniliform cross sections. Interzooidal pores are small in diameter (about .002 to .003 mm) and are seen infre- quently (Text-fig. 14D). Zooecial chambers have elliptical to sub- elliptical cross sections. Diaphragms — Intermediate diaphragms are common in zooecia subjacent to overgrowths. The diaphragms are generally seen about 0.1 to 0.3 mm aboral to the aperture (PI. 32, figs. 1f, g; Pl. 33, figs. 2a-c; Pl. 34, figs. la, b, 2b). Less commonly, diaphragms are scat- tered throughout the overgrowth unit (PI. 34, fig. 2b). The dia- phragms generally are thin (about .003 to .006 mm) and nearly planar, and show slight aboral flexure at the juncture with the zooecial wall before merging continuously with the zooecial lining. Basal diaphragms were seen rarely, occurring just subjacent to the boundary of zoarial growth phases in zooecia whose chambers are uninterrupted by basal layers. Brood chambers — Brood chambers occur commonly. The chambers are wide but shallow (see Table 19) and lenticular with slightly convex proximal and distal surfaces. The chambers occur in the more proximal portions of a single overgrowth unit (PI. 32, figs. le, f; Pl. 34, fig. 2b). Most subjacent zooecia are closed by the brood chamber floor; a few pass through the chamber to the roof. These zooecia are commonly thin-walled and continuous with thin-walled, septate partitions (Pl. 33, fig. 1). The partitions radiate laterally from the open central portion of the chamber. The brood chamber roof is thick (about .03 to .04 mm); pores are about .01 mm in diameter. BULLETIN 291 118 ‘UOTIIIS ASIDASULI} UI paInseauw ‘9UOZOXa IaUU] :Z sUOZOX| ‘uo1jda8 [eUaSue} Ul poInseaw ‘au0zZOXxd 13}NGQ :] IU0ZOX| SIOJIUIT|[IW UT UGXINN-USSD-4O9Z UGXW-USS-YQ2Z UQXWN-USs8)-YO9Z UQXIN-USs)-YOIZ Z 9U0zZ0Xy - [8199007 US809Z/UD FdPZ YL-LM9PZPO UGX N-US8)D-4O9Z UQXIN-IS8)-YO9Z c T 0s 0s 1d eT OT V7 c T 0s 0s 0€ Z0° 90° £0° or Z T 0s 0s 9¢ Z0° 80° +0° tT e 0s 0S 0 4 T Z SZ SL ce TIO +£0° $T0° 020° T c SZ SZ el c cl OT 31 T c SZ SZ SC: c0" 90° ZO" i) I Z SL SZ 1x4 c0° 80° £0° IT’ apod ‘sedg IZN 9ZN N “ACO aS #X UGXINN-USSD-499Z UCXIW-USSQ-YO2Z I 9u0zoxg - [8199007 IayoeieyD (SSQAHd10D) FYOdOLdAYD FAYOdONALIAH AO SLNAWAYNASVAW AO AUVWWAS TVOILLSILVLS 6l ATAVL mg NYE CERIOPORID CycLosToMEs (Bryozoa): (81) TI- ‘($Z) 8- “(9T) 9- “(1T) S-Z8EZ INS SedAjodoy, + (¢Z) addjoxoa]eseg ‘(€Z) BIT AN edAiojaT “+ SIL ag edAj03}007T “¢ ($2) 2-L8EZ “207 WNSN edéjodoy, * (SZ) B8IT a adAjo}99T °Z ($2) 2-L8E% 907 WNSNA edAjodoy, (0S) BIT an 2d4}0}097T “1 ado0o NAWIOddS OL ATM ‘U0I}N9S ISIDASULI} UI PainseaWl ‘sUuOZOXxa I9UUT +7 9UOZOX| SIOJIWIT[ [IU UT» £ I I SZ10° uqxW-USso-1dF AOA 3 I Or II +00" +£0" 820" 0+0° YL-FauOId 3 I Z + wd-4old 3 I Z ST 8T WA-YOld Joquieyg poorg + 9 811 SII 0€ +0° ae Z0" 97° ydyouq-qiuy useiydeid g 0s 0s 0 aT US8D2Z /UD-1d PZ Z I 0s 0s tt +10" Z£0° LT0° 0L0° YL-IM9ZPO apoj92dS_ = IZN OOZN N ‘A‘O *S #X «WO ToyoBIeYyO (SSQHUGTOD) FYOdOLdAYD FYOdMONALAH AO SLNAWAYASVAW AO AUVWWAS TVOLLSILVLS 120 BULLETIN 291 TABLE 20 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN HETEROPORA CRYPTOPORA (GOLDFUSS) Cire: Ellip. Ov. Pyr. Polyg. Triang. Irreg. Outer Exozone Regular = 35 9 Sub 7 16 1 2 Irregular 1 Inner Exozone Regular 1 26 3 Sub 9 10 1 Irregular Discussion. — Von Hagenow (1851, p. 53) returned the species H. cryptopora to Ceriopora and singled out only one specimen of the syntype suite to bear the name C. cryptopora. This restriction was made in his synonymy of C. cryptopora by the citation, “Certopora cryptopora Goldfuss. Th., Petr. I, p. 33, Taf. X, fig. 3, a (nicht b-d).”, and his statement in the remarks following that: “Mir ist nur dieses eine Exemplar in Bonnenser Museum, von Maastricht bekannt.” Von Hagenow then assigned the remaining figured speci- mens (Goldfuss, 1826, pl. 10, figs. b, c, d) to other species of Hetero- pora. Gregory (1909, pp. 188-9) carefully reviewed Von Hagenow’s analysis of the specimens from the syntype suite of C. cryptopora and remarked that it was “simplest to restore Goldfuss’ conception of this species”. He considered that all of the specimens of the syn- type suite illustrated by Goldfuss (pl. 10, figs. 3a-d) as C. cryptopora were conspecific; and in doing so, he consistently referred to the specimen illustrated by Goldfuss (pl. 10, fig. 3a) as the type. Gregory’s arguments were, in turn, followed by Borg, 1933p: 283). Borg stated that: I think, however, that von Hagenow’s ideas as to the suitable limits of a species were a little exaggerated, and I am not convinced that the specimens figured by Goldfuss (of. cit.) in his Pl. X, figs. 3a, 3b, and 3c, did not all belong to one and the same species. The differences existing between them seem to me to be easily explainable by the fact that they obviously represent different portions of three zoaria. CERIOPORID CycLosToMEs (Bryozoa): NYE VAI 751A 75 B ~> > oO Oo = (= 250 250 oT o z 2 PS 25 mm 4a 42 nes 1.0 2.0 ZcCh-CsSn-NMxDn ZcCh-CsSn-Mx Dn ZcCh-CsSn-NMxDn 751C ~> is 100 is o =o x75 4 oO © = = = SO 25 iS = = 25 pe} oO mm af 4p NS) "45" 5)6 CdZcWI!I-Th ZdPr-Cn/ZcCsSn Text-figure 14 A-D. Histograms and cumulative curve from the lectotype and one topotype of Heteropora cryptopora (Goldfuss). A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness. D. Count of interzooidal pores per zooecial cross section. 122 BULLETIN 291 Designation of the lectotype here follows the previous restric- tion of Von Hagenow (1851) and later followed by Gregory (1909) and Borg (1933). Remarks on morphology. — Diaphragms were interpreted to be intermediate, rather than basal, because a few show slight aboral flexure at the juncture with the zooecial wall. There is apparently a relationship between diaphragm emplace- ment and zoarial growth cycles. Intermediate diaphragms commonly occur singly in a zooecium and are commonly emplaced a short distance proximal to the terminal surface of an overgrowth unit. Generally, only zooecia with chambers continuous orally into the superjacent growth unit lack diaphragms. The taxonomic and phylogenetic significance of the zoarial mode of growth exhibited by H. cryptopora is at present unknown. Description and illustrations of a number of species presently thought to be assignable to many diverse genera and families suggests that repetitious addition of intrazoarial overgrowths may be a relatively widespread mode of zoarial growth in cyclostome bryozoans, and some trepostomes as well (Boardman, 1960, pp. 39-40, 57-58, pl. 7, fig. 4, for Leptotrypella multitecta Boardman, a Devonian trepo- stome). Below is a partial listing of a few species and illustrations suggesting the occurrence of this mode of growth in Cyclostome Bryozoa: In Canu and Bassler, 1926 — Ceriopora falax, text-fig. 13, p. 28; Diplocava globosa, text-fig. 38, p. 74; Multicrescis lamellosa, text- fig. 2, p. 14; Multigalea canu, text-fig. 31, p. 62. In Gregory, 1909 — Multicrescis tuwberosa, text-fig. 54, p. 207; Radiopora neocomiensis, text-fig. 74, p. 285; Reptomulticava fungi- formis, text-fig. 39, p. 136. In this study, incremental addition of intrazoarial overgrowths was seen to play a major role in the zoarial growth of Diplocava incondita and, to a lesser extent, Ceriopora micropora. Genus LEIOSOECIA Canu and Bassler, 1920 Type species: Multicrescis parvicella Gabb and Horn, 1861, by original designation and monotypy, Canu and Bassler (1920, p. 823). 1920. Pars Leiosoecia Canu, and Bassler, U.S. Nat. Mus., Bull. 106, p. 823. 1922. Pars Leiosoecia Canu and Bassler, Canu, and Bassler, Wis. Nat. Mus., Proc:; vol: 61; p. 99: CERIOPORID CycLosToMEs (Bryozoa): NYE 125 1953. Pars Leiosoecia Canu and Bassler, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G72. Tentative diagnosis. —Zoaria branched; branches with well- differentiated coaxial exozones and endozones. Distally growing branches commonly intersecting and anastomosing. Zooecia dimorphic. In exozone, zooecial walls distinctly laminate. In profile sec- tion, laminae wavy, sometimes crenulate, forming irregular V- to U- shapes convex orally. Zooecial lining present. Intermediate diaphragms occur occasionally in outer exozone. Taxa included. — Only the type species, L. parvicella, is here included. Internal characters of other species assigned to Leiosoecta are unknown to me. Discussion.—Canu and Bassler (1920, p. 823; 1922, p. 99) based their definition on secondary specimens of the type species. These secondary specimens are not considered to be congeneric with the lectotype. The original definition by Canu and Bassler (1922, p. 99) was extremely brief: “Zoarium consisting of cylindrical tubes and regular, parietal mesopores”, and could apply to many tubular bryozoans. In 1953, Bassler added the description of the brood chamber to the definition and figured it. This addendum was also based on secondary specimens identified with the type species by Bassler but not considered here to be congeneric with Letosoecta. Leiosoecia, in its simplicity of appearance and mode of growth, shows close morphologic similarity to Tetrocycloecia Canu. The morphologic differences between them are considerably less than differences between the other genera studied. In Leiosoecia, however, dimorphism is poorly expressed in zooecial dimensions, and inter- mediate diaphragms are occasionally seen. Leiosoecia parvicella (Gabb and Horn), 1861 Pl. 36, figs. la-h 1861. Multicrescis parvicella Gabb, and Horn, Acad. Nat. Sci. Philadelphia, Proc., vol. 12, p. 367. 1861. Multicrescis parvicella Gabb and Horn, Gabb, Acad. Nat. Sci. Phila- delphia, Jour., ser. 2, vol. 4, p. 401, pl. 69, figs. 36-38. 1861. Multicrescis parvicella Gabb and Horn, Gabb, and Horn, Acad. Nat. Sci. Philadelphia, Jour., ser. 2, vol. 5, p. 178, pl. 21, fig. 70. 1905. Multicrescis parvicella Gabb and Horn, Johnson, Acad. Nat. Sci. Phila- delphia, Proc., vol. 57, p. 5. 1907. Non Heteropora parvicella (Gabb and Horn), Ulrich, and Bassler, Geol. Sur. New Jersey, Paleont. Ser., vol. 4, p. 327, pl. 23, figs. 1-2. 1920. Non Leiosoecia parvicella (Gabb and Horn), Canu, and Bassler, U. S. Nat. Mus., Bull. 106, p. 823, text figs. 273A-F. 124 BULLETIN 291 1922. Non Leiosoecia parvicella (Gabb and Horn), Canu, and Bassler, U-S. Nat. Mus., Proc., vol. 61, p. 100. Type. — ANSP 31261 is designed as the lectotype. The lecto- type is the only syntype known to be preserved. The specimen, ANSP 31261, compares favorably in zoarial form and surface ap- pearance to Gabb and Horn’s illustration (Gabb and Horn, 1861, pl. 69, figs. 36-38) and is apparently the specimen referred to by John- son (1905) as the figured specimen. ANSP 31261 is labeled “Type”. Type locality and horizon. — The label with the lectotype bears the inscription, “Timber Creek, N.J.”, one of the two localities originally listed by Gabb and Horn. The specimen probably comes from the Vincentown Formation (pers. comm. H. Richards, 1967) of Paleocene age. The age and distribution data cited for L. parvicella is here considered questionable. FE. O. Ulrich, R. S. Bassler, and others gathered extensive collections of bryozoans from a number of localities in New Jersey and Delaware, including Timber Creek, the cited type locality. These collections, now housed by the National Museum of Natural History and the United States Geological Survey in Washington, D.C., were searched for specimens bearing resem- blance to the lectotype; none was found. Material studied. —The lectotype was kindly loaned to the author by Horace Richards, Academy of Natural Sciences of Phila- delphia. Most of the original zoarial fragment remained intact after six thin sections and four acetate peels were made. Duplicate peels are preserved in the National Museum of Natural History collection and the author’s collection. Description of the lectotype.— This description is based solely on the lectotype, the only specimen of L. parvicella known. This description includes assessment of nongenetic variation within this colony. No assessment of genetic or other interzoarial variation with L. parvicella is implied in this description. Growth habit — Branches are roughly cylindrical. The zone of zooecial bending is relatively broad and begins commonly deep in the endozone (PI. 36, fig. le). Zooecial growth axes are moderately undulatory in both endozone and exozone (PI. 36, figs. 1d, e). Endozone — The zooecial walls are thin and commonly parallel- CEeRIoporID CycLosToMEs (Bryozoa): Nye 125 sided, sometimes with a small variation in thickness giving sub- moniliform cross sections (PI. 36, fig. le). Zooecial chambers are polygonal to subpolygonal in cross section. Interzooidal pores are rare. Zooecial walls are homogeneous to subgranular with thin, dark zooecial linings. Exozone — Large dimorphs are commonly surrounded by small dimorphs, less commonly are directly adjacent to another large dimorph (PI. 36, fig. 1f). Zooecial walls of large and small polymorphs are similar in appearance. Both commonly show moderate asymmetry in thick- ness across the zooecial boundary zone, and both have moniliform profiles due, in part, to the flare of interzooidal pores and to the longitudinal variation in wall thickness (PI. 36, fig. 1g). Walls of large polymorphs, however, generally appear to be more parallel- sided in profile. Interzooidal pores were rarely observed (Text-fig. 15C). Diaphragms — Intermediate diaphragms were observed ap- proximately .02 to .05 mm aboral to aperture in both large and small dimorphs (PI. 36, fig. 1g). Diaphragms are nearly planar and flex aborally at the juncture with the zooecial wall to merge with the zooecial lining. TABLE 22 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN THE LECTOTYPE OF LEIOSOECIA PARVICELLA (GABB AND HORN) Circ. Ellip. Ov. Pyr. Polyg. Triang. Irreg. Exozone Regular 7 44 4 1 Sub if 26 3 Irregular 2 + 2 Endozone Regular 7 Sub 2 1 11 2 Irregular 2 BULLETIN 291 126 SI9JIWT [IW UTy UQXINN-US8D0-4O9Z I if 9b 9+ 8T A Cal OT ST UQXW-USsp-YOXZ T T oF 9+ 8T c0° or 90° A UGXINN-USSD-YDIPZ T T oF oF LI c0° oh LO’ 07 UQXIN-USsO-YOIZ sydiowAjog as81eq - 9u0z0xy - [e199007 T T £s £¢ 0 aT » USSD9Z/UD-1d PZ T T £¢ £S 9+ 20° +0° TO’ 0) & YL-ILM2ZPO UQXWN-USSD-YD9Z T T £$ €£$ 61 g vT OT Ig UqX*W-USs)-YD92Z T T €$ £s 972 c0° 90° 0° Or UGX N-USSD-YDIZ T if £S £S £7 c0° 80° £0° (Ae UQXIN-USsD-YOIZ sydiowAjog [[BWsg - 3u0Z0xy - [8199007 T I t OG 83°C UQXW-USSD-1g [Bl1e07 epoyd ‘vadg IZN PN N ANNO) #S aX Pap: MO) ToJOBIe YS) (NYOH GNV AEVD) PITAIIANFd FIDAOSOIAT AO SLNAWAYASVAW AO AUVWWAS TIVOLLSILV.LS 1? ATAVL 127 CeRIopoRID CycLosToMeEs (Bryozoa): NYE adqd00 NAWIOdds OL AAU ‘I9ZTE USNV ‘2d4}0399T “T SIOJIWIT][IU UL y I Lee se SZ 0 | UgsQ2Z /UD-1gPZ I 1 §2 6 02 100" £00" $00 O10" YL-LM2ZPO UX N-USS9-4Q9Z I 1 sz $2 $1 z’ oT Or 91 UAXW-USs)-4Q9Z I 1 sz $z 0g £0° or +0 +1 UCXWN-USSQ-492Z I 1 sz $2 lz +0' ae 90° 61" UqXIW-USsQ-492Z auoZzOpUuy - [B199007 I Tce 66 0 Z USsQ2Z /UD-1qPZ I Tomi Oor ar ZO" +0" 10° or YL-IM2ZPO UGX N-USS9-4D2Z I 1 001 oor 61 2° £1 Or 12 UdXW-Ugs)-499Z L T Oot OOT Se £0° 80° c0° rT UGXWN-YUSSD-YDIPZ I T 00m oormee os 60° or £0° 02" uqxXW-Ugs)-499Zz sydiow jog |] VY - 2u0z0xg - [e199007 I Tae St 0 7 USsQ2Z /YD-14PZ I De oe oF Zb Z0" +0 10° or Y.L-IM2ZPO ‘apog‘20dg =IZN OZN N ais +X «XO yayORIEYO (NYOH GNV EEVD) PITZDIANFd FIDAOSOIAT AO SLNAWAYNSVAW AO AUVWWAS TIVOLLSILV.LS 128 BULLETIN 291 751A 75 B ~ ~ oO (6) fox (os 250 250 oO oOo 2 S Nh oO Nh oO mm 4 a2. a8) 1.0 2.0 ZcCh-CsSn-NMxDn ZcCh-CsSn-Mx Dn ZcCh-CsSn-NMxDn On N (e) On NO oOo cumulative % i 23? 4 ores ZaPr-Cn/ZeCssn Text-figure 15 A-C. Histograms and cumulative curve from the lectotype of Letosoecia parvicella (Gabb and Horn). A. Normal to maximum cross- sectional dimension of a zooecial chamber. B. Ratio of the maximum cross- sectional dimension of a zooecial chamber to the normal to maximum cross- sectional dimension of a zooecial chamber. C. Count of interzooidal pores per zooecial cross section. CERIOPORID CycLosToMEs (Bryozoa): NYE 129 Discussion. — The concept of Leiosoecia, as developed in the descriptions and illustrations of Ulrich and Bassler in Weller (1907), Canu and Bassler (1920, 1922) and Bassler (1953), was based on observations of secondary specimens assigned to Letosoecia parvt- cella. Thin sections prepared by R. S. Bassler and published illus- trations were examined. In addition, a few other secondary speci- mens from the Paleocene, Vincentown formation, Blackwoodstown, and Vincentown, New Jersey, identified as L. parvicella by Bassler, were thin-sectioned and examined. The secondary specimens (Canu and Bassler, 1920, p. 824, text-figs. 273A-C) are branched, and have coaxial endozones and exozones. The branches, however, do not anastomose, and are small and delicate in appearance in contrast to the robust anastomosing branches of L. parvicella. Zooecia are dimorphic in the secondary specimens (Canu and Bassler, 1920,"p: 824, text-figs. 273D-F). The small polymorphs are irregular, nearly sinuous in growth. The large polymorphs commonly exhibit a cylindrical sheath of homogeneous calcareous tissue around the zooecial chamber similar to that seen in Parleiosoecia. There are thick deposits of interzooidal calcareous tissue in the exozone simi- lar in appearance to that commonly seen in hornerids. Intermediate diaphragms are numerous. Comparison of this preliminary characterization of the secondary specimens to that of the lectotype of L. parvicella reveals significant morphologic differences. These differences are considered to be at least as significant as those which are used to differentiate other genera treated here. Thus, the secondary specimens are not con- sidered here to be congeneric with the lectotype of L. parvicella. Genus PARLEIOSOECIA Canu and Bassler, 1920 Type species: Parleiosoecia jacksonica Canu and Bassler, 1920, by original designation and monotypy. 1920. Parleiosoecia Canu, and Bassler, U.S. Nat. Mus., Bull. 106, p. 824. 1953. Parleiosoecia Canu and Bassler, Bassler, Treatise on Invertebrate Pale- ontology, Part G, Bryozoa, p. G72. Tentative diagnosis. — Zoaria thin, encrusting expansions, or branched. Branches have strongly differentiated exozones and endo- zones coaxial with series of hemispherical chambers. Chambers 130 BULLETIN 291 formed by extensions of basal layer above substrate. Basal layer laminate; laminae inclined proximally oblique. Intersection and anastomosis of distally growing branches common. Zooecia dimorphic. In endozone, laminate tissue lines zooecial chamber entirely, commonly thickest at proximal tip and deposited directly upon basal layer. In exozone, cortex of large dimorphs composed of light-colored, homogeneous-appearing calcite. Walls of small dimorphs distinctly laminate. Laminae commonly broadly arched, orally convex, and continuous across boundaries with adjacent small dimorphs. V-shaped patterns occasionally seen. When adjacent to large polymorphs, laminae abut cortex of large poly- morphs at relatively high angle. Terminal, intermediate, and basal diaphragms present. Taxa included. — Monotypic for the type species. Discussion. — P. jacksonica achieves erect branching habit by an unusual modification of the encrusting mode of growth. Locally, the basal layer grows above the encrusted substrate and is extended as a series of hemispherical chambers. The structure produced forms the axial support of a distally growing branch which, in turn, branches to produce a zoarium with ramose growth habit. Branching at short intervals and anastomosis of distally growing branches as seen in zoarial fragments, suggest that the colony so produced was at least moderately large and densely branched in the proximal por- tions of the colony. Large polymorphs bud from the latero-distal portion of each hemisphere, but apparently never bud from, or over- grow, the distal tip of the branch, 7.¢., the distal part of the hemi- spheric-axial chamber composed of the basal layer. The basal layer lacks pores and is structurally different from the calcareous tissue of zooecial walls. Also, the basal layer is often separated from the recumbent zooecial walls by a dark line. The structural characteristics and the size and configuration of the chambers make it difficult to believe that they are zooecial poly- morphs. Similar-appearing axial structures were described by Pergens (1890, p. 318, text-fig. 11) in Cavaria von Hagenow, and in Sezn- laterotubigera d’Orbigny. Pergens stated that, “leur réle est inconnu; CerrIoporip CycLosToMeEs (Bryozoa): NYE 131 peut-etre servent-elles a la reproduction” (their function is unknown; perhaps they serve for reproduction). Such a function seems unlikely in Parleiosoecia gacksonica. The only resemblance to brood chambers in other cyclostome species is the formation of a relatively large chamber. The axial chambers differ in position and in structure. Communication pores to adjacent zooecia were not observed, and there are no subjacent zooecia. In P. jacksonica, the wall of the axial chamber is an extension of the basal layer, but brood chamber walls are homologous in structure with zooecial walls and not with basal layers. Brood chamber roofs are commonly (but not invari- ably) porous. Only a single, central opening was occasionally ob- served in the axial chambers of P. gacksonica. Finally, hollow struc- tures with the typical appearance of brood chambers have been identified in several specimens of P. jacksonica; these are located in the exozone and are unrelated to the axial chambers. The formation of hollow branches by budding of zooecia from a cylindrical-appearing basal layer is known in several cyclostome species such as Seminodicrescis nodosa d@Orbigny, Cavaria ramosa von Hagenow (Gregory, 1899, text-fig. 54, p. 400) and Spiropora macropora d’Orbigny (Pergens, 1890, text-fig. 11, p. 318). Often this may be explained by encrustation on a previously existing struc- ture. In Recent specimens of Denstpora corrugata Macgillivray, bits of a tubular, woody, marine plant are often preserved which the bryozoan encrusted. Hamm (1881, p. 25) suggested that axial cham- bers in Cavaria pustulosa were caused by encrustation on a soft stem. Gregory (1899, p. 399) stated that this conclusion was “un- tenable”, but suggested no alternatives. The nature of encrustation in P. jacksonica can, at present, be inferred only from negative evidence. Remains that could be inter- preted to be a substrate organism or structure have not been recog- nized in the axial chambers. The outer surface of the basal layer is smooth to rugose, but impressions suggesting encrustation on a sub- strate organism were not observed. Partitioning of the axial hollow by the basal layer suggests that the cavity was essentially empty when the partition was emplaced. The evidence, as presently under- stood, suggests that the axial chambers did not encrust a previously existing structure formed by another organism, but that the secreting 132 BULLETIN 291 epithelium built its own substrate, the basal layer, as growth con- tinued distally. Parleiosoecia jacksonica Canu and Bassler, 1920 Pls 3%, figs alta-cs Pl; 38, figs. 1, 2, 3a, b, 4a-c; Pl..39, figs. 1, 2a-c, 3, 4, 5; Pl. 40; fiss#ia-t92 1920. Parleiosoecia jacksonica Canu and Bassler, U.S. Not. Mus., Bull. 106, pp. 824-5, text-fig. 208e, p. 646; text-fig. 274a-c, p. 825; pl. 148, figs. 1-13. 1953. Parleiosoecia jacksonica Canu and Bassler, Bassler, Treatise on Inverte- brate Paleontology, Part G, Bryozoa, text-fig. 37, figs. 4a, b, p. G72. Type. — USNM Loc. 2933B-1 is here designated as the lecto- type. [his specimen is figured here in Plate 37, figures la-g, and was figured by Canu and Bassler, 1920 (pl. 148, fig. 2). Type locahty and horizon.— The lectotype was probably col- lected from the Eocene, Jacksonian, Eutaw Springs, South Carolina. The original label with the specimen bears only the word Eutaw. Material studied. — Five thin-sections and 11 acetate peels were made from the lectotype. Thin-sections and acetate peels were made from two of the paralectotypes: USNM 65447-1 figured by Canu and Bassler in Pl. 148, fig. 1; and here, Pl. 27, figs. 2a-c; USNM 65449 figured by Canu and Bassler in text-fig. 274A, p. 824, pl. 148, fig. 6, and here, PI. 28, figs. la-f. In addition, thin-sections and acetate peels were made from 31 specimens from Eutaw Springs, S.C., USNM Loc. 2933A. Dupli- cate acetate peels of all specimens except USNM Loc. 2933A-1-5 are in the author’s collection. There were many inconsistencies between data as given on original labels, as cited in Canu and Bassler (1920), and as cited in catalogue entries for the specimens. Following is a list of the syntype specimens with catalogue numbers and locality data as given on the original labels. Also included is a listing of illustrations be- lieved to be made from a particular specimen, arrived at by direct comparison of the specimen with illustration. The labels on Bassler’s thin-sections have plate and figure citations on the original label. It was not possible, however, to identify each section with the cited illustration. In such instances, the citation as originally cited is listed with a question mark. Cat. No. 65446; 5 thin-sections, Jacksonian, Rich Hill, Georgia. -1 ? Pl. 148, fig. 9 -2 ? Pl. 148, fig. 10 Cerroporip CycLosTomes (Bryozoa): Nye 133 -3 ? Pl. 148, fig. 11 4 PI. 148, fig. 12 -5 Pl. 148, fig. 13 Cat. No. 65447: 2 specimens, Middle Jacksonian, Santee River, 3 miles above Lenuds Ferry, S.C. “1° Pl: 148, fig. 1 =2' Pl. 148, figs. 4, 5. Cat. No. 65449: 1 specimen, Eocene (Jacksonian Middle), 18 miles west of Wrightsville, Johnson Co., Ga., text fig. 274A, p. 824, pl. 148, fig. 6. No Cat. No.: 4 specimens apparently not catalogued at the same time as the other specimens. Eocene, Middle Jacksonian, Eutaw Springs, S.C. = P 148; fig: 2 Remainder unfigured. Description. — Mode of growth — Branches are commonly cylindrical, occa- sionally frondose. The distal-most portion of the axial chamber is sometimes open, with the basal layer flexing distally to form a lip around the opening (PI. 38, fig. 4b; Pl. 39, fig. 4). A narrow zone of zooecial bending separates the” well-differentiated exozone and endozone (PI. 38, fig. 3a). Endozone — The walls of large polymorphs are thin and parallel-sided. Large polymorphs are slightly undulatory in their growth, are inclined at a low angle to the branch axis (commonly less than 30°), and are long (approximately 1.5 mm from proximal tip to zooecial bend). Large polymorphs but in ranges parallel to a branch axis. Zooecia in adjacent ranges are budded alternately. The overlap of adjacent recumbent zooecia is commonly ac- complished by the interwedging of prismatic zooecia which have regular polygonal cross sections (Pl. 39, fig. 5). Sinus and keel ac- commodation is seen less commonly. The budding pattern is ex- pressed in the exozone by a more-or-less regular, rhombic distribu- tion pattern of large dimorphs (PI. 39, fig. 2a). Exozone — Large dimorphs have thin, parallel-sided walls which show little change in thickness throughout the exozone. The walls project slightly above the zoarial surface. Zooecial chambers of 134 BULLETIN 291 large dimorphs are commonly elliptical to subcircular in cross sec- tion. Interzooidal pores are seen rarely. The longitudinal profiles of the zooecial walls in small poly- morphs are somewhat variable in appearance. The walls sometimes show little longitudinal variation in thickness, sometimes show grad- ual increase orally (PI. 40, fig. 1f), or show longitudinal, commonly annular, variation in thickness. The walls are commonly thickened subsymmetrically across zooecial boundary zones, but circular or clavate monilar cross sections are often observed. Zooecial chambers have subcircular to elliptical cross sections. Diaphragms — Terminal diaphragms sometimes occur. General- ly, the diaphragms are slightly (about .055 mm) subapertural in position. The diaphragms have planar oral surfaces and orally convex aboral surfaces. They have short aborally flexed abutments. Intermediate diaphragms are common in occurrence but fre- quently show large variability from zoarium to zoarium. Generally, no more than one diaphragm occurs in a single zooecium. Neighbor- ing zooecia commonly have diaphragms in similar positions (PI. 38, figs. 3a, 3b; Pl. 39, fig. 2b). The diaphragms are thin and planar to slightly convex orally, and they flex aborally at the juncture with the wall to either merge with the zooecial lining, or to form a short, thin abutment distinct from the zooecial lining. Basal diaphragms were rarely observed, and occur in both large and small dimorphs. The diaphragms are thick, with slightly wavy laminae about parallel to the diaphragm surface. The diaphragms are orally flexed at the juncture with the zooecial wall. Brood chambers — Brood chambers occur in the middle and outer exozone, and are lenticular to subconical with a planar to domal roof (PI. 40, figs. la-e). The walls of zooecia subjacent to brood chambers often become thin-walled (to about .01 mm) as much as .1 mm proximally from the brood chamber (PI. 40, fig. 1d). The brood chamber floor is thin (less than .01 mm). Most zooecia are sealed at the floor of the brood chamber, but a few large polymorphs pass continuously through the chamber to the roof (PI. 40, figs. 1b, c, 2). Intra-brood chamber zooecia have thin, parallel-sided walls (PI. 40, figs. 1c, e). The intra-chamber zooecia are often continuous with vertical septate partitions distributed radially away from the center 135 SIOJIWT| [TU UT» UQXINN-USSD-YO2Z $ c 02 0c 9 v TT OT cl OGSIN=US§O 192Z ra) $ c 02 02 9 T0° 60° 80° or UQXxINN-YSS)-4O9Z Ss = Pd $ c 0c 02 Or 10° or 60° cr UGAWs tS Os O2Z oe auo0zoxy - sydiowAlog adie - [e109007 ow 3S v £ SL SL bE £00° cT0° 900° 10° YL-IMPZPO S suozopuy - sydiowdjog [[V - [B199007 cs £ £ O1T OrT 0 at ug8)9Z/UD-1dPZ S £ £ OTT OIL OF 10° £0° 600° 690° YL-LMPZPO a UCXIAN-USSO-Y2Z 2 £ £ OTT OTT LI c raat OT vc UGSINsUSEDs199Z 2 £ £ OTL OTT Sf c0° $0° c0° Or UCXxINN-¥USS0-YO9Z o £ £ OT Ort ee 20° 90° £0° eT BG=INUS OMI Ss au0zoxg - sydiowAjog [|[Vy - [B199007 a Z II II z 200° 010" 800° Z10° YL-AATISE Z =, T L L £6 v + c vi UGS ASSO MOV - I L L 61 v 4 LT LO VGSINGGSs Osta i [B11v0Z O apoD ‘vad IZN ZN N “AO *S #X UO) YaTSSVa UNV ONVO FPOINOSNOFL FIDJOSOIAIAFd AO SLNANAYASVAW JO AUVWINDS IVOLLSILVLS ef ATAVL BuLLeETIN 291 136 T-6btS9 WNSOA adAoajeieg °g "(OL) 9-VEE6C “20T WNSQ ‘12[8seg Aq parfynuap! adAjodoy, *Z *(6) 8- ‘(Tb) 9-VEE6Z 207 WNSA “121sseq Aq palynuap! sedAjodoy puke *(€b) I-A££62 90T WNSA AdAio127T °9 (OL) 9-VEE6Z 20T WNSA ‘22I8segq Aq palynuepr addjodoy pue (OT) T-AEe6z 0T WNNSN 244303997 *s adoOO NAWIOddS OL ATM "($Z) II- ‘(sz) ST- ‘($Z) €2-VEE67 “20T WNSOA “Jatsseg Aq patynuap! sadAjodoy, “> ‘(O1) 8- ‘(0S) 9-VEE6C 207 WNSO ‘lajsseg Aq pel}uap! saddjodo} pue ‘(0¢) I-g£f6c 207T IWNSO ‘9d.4}0}D9T *¢ "() €2- §(9) TI-VEE6z 207 WNSDO “aisseg 4q palynuapr sadAjodoy, *Z "VEE6Z “90T IWWNSD ‘t2]sseg Aq parynuap! SadAjodoy 9 puke I-G£e6c “907 WNS1) 2d4I02997 “1 SIOJIUINT|[IW UT, 8 I Ciemnou, eec00: 110° 100" +10 1d-FUNOI ad 8 E s s1 +00" £60" 620" 860° UL-FUYO4d 8 if I aT WM-YyOlg laquivyy poolg 9 € £6 £6 0 it USsD9Z/4D-1d PZ UQXWN-4S80-4YO9Z 9 £ £6 £6 91 c aa OT +2 UX -USSD-YDIZ 9 £ £6 £6 it 10° +0" Z0" 90° UGXIWN-USS9-499Z : : ‘- i i ze = oa . auo0zoxy - sydiowAjog A Eues eee s Z 02 02 0 USs99Z/4D-1dPZ S Z 0z 02 9¢ 600° 970° Z10° 6+0" U.L-IM2ZPO IZN ZN N eee) Ss *X * TO JayoeIeyy apoD ‘vadg UATISSVA GNV ANVO PIINOSNOFS FPIDAOSOIATA Fd AO SLNANAANSVAW AO AUVWIWAS TVOLLSILV.LS Cer1oporip CycLosTomMeEs (Bryozoa): NYE 137 754A 75 B > ~> oO oO c = 250 250 oO lox g : 25 25 mm 4d 2. 7S 1.0 2.0 ZcCh-CsSn-NMxDn ZcCh-CsSn-Mx Dn ZcCh-CsSn-NMxDn 754C > 100 oO ce ®o = 50 75 D oO ® > _ a 50 25 = a = 25 = | oO mm Al a2 it 2) ona 52.0 CdZcWI!-Th ZdPr-Cn/ZcCsSn Text-figure 16 A-D. Histograms and cumulative curve from the lectotype and two topotypes of Parleiosoecia jacksonica Canu and Bassler. A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness. D. Count of interzooidal pores per zooecial cross section. 138 BuLLeTIN 291 of the brood chamber, but leave the central area open. The roof is thick (about .03 mm) and porous (pores up to .01 mm in diameter). When abandoned, the brood chamber is submerged by a basal layer extending from lateral zooecia, and new zooecia bud from the basal layer. TABLE 24 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN PARLEIOSOECIA JACKSONICA CANU AND BASSLER Gire) ) SE llip: Ov. Pyr. Polyg. Triang. Irreg. Large Polymorphs - Exozone Regular 3 8 Sub 7 2 Irregular Small Polymorphs - Exozone Regular 2 50 8 Sub 3 20 + Irregular 5 Remarks on mode of growth. — Boardman and Utgaard (1966, pp. 1033, 1036, text-fig. 1, p. 1083) reconstructed the three-dimen- sional shapes of zooecia in several encrusting Paleozoic bryozoans. The recumbent endozonal portions of zooecia commonly developed an interlocking sinus and keel configuration. This configuration was apparently related to the utilization of available space, and was re- flected in orderly budding patterns and in the regular arrangement of zooecial apertures as seen at the zoarial surface. Boardman and Utgaard believed that this configuration might be widespread in encrusting tubular bryozoans. In P. jacksonca, the appearance of endozonal zooecia as seen in transverse section, e.g., first row triangular to hemispherical, second row submushroom shape, is generally comparable to the cross sectional shapes of zooecia as described and illustrated by Boardman and Utgaard, suggesting that the recumbent, endozonal portion of zooecia also have an interlocking sinus and keel configura- tion. In P. jacksonica, this pattern is modified because the large polymorphs are oriented radially from, and are recumbent upon, a nearly spherical surface (the axial chamber) rather than a nearly flat surface as seen in the encrusting, sheetlike colonies examined CERIOPORID CycLosToMEs (Bryozoa): NYE 139 by Boardman and Utgaard. Accommodation to this spherical sur- face may, in part, explain the less regular polygonal shapes seen in P. jacksonica (Pl. 39, fig. 5). Genus REPTONODICAVA d’Orbigny, 1854 Type species: Ceriopora globosa Michelin, 1846, by subsequent designation, Bassler (1935, p. 186). 1846. Pars Ceriopora Goldfuss, Michelin, Iconographie Zoophytologique, Description par Localités et Terrains des Polypiers Fossiles de France et Pays Environnants, p. 246. 1854. Reptonodicava d’Orbigny, Terrain Crétacé Bryozoaires: Paléontologie Francaise: Description des Animaux Invertébrés, vol. 5, p. 1014. 1896. Pars Ceriopora Goldfuss, Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Jurassic Bryozoa, p. 195. 1909. Pars Ceriocava d’Orbigny, Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Cre- taceous Bryozoa, vol. 2, p. 127. 1920. Pars Ceriopora Goldfuss, Canu, and Bassler, U.S. Nat. Mus., Bull. 106, p. 678. 1935. Reptonodicava d’Orbigny, Bassler, Fossilium Catalogus, I, Pars 67, Bryozoa, p. 186. 1953. Pars Ceriopora Goldfuss, Bassler, Treatise on Invertebrate Paleontology: Part G, Bryozoa, p. G67. Tentative diagnosis. — Zoaria globose, hemispherical to cylin- drical; sometimes with bulbous to branchlike outgrowths or having less regular, massive shapes. Zoarial surface smooth or uneven, com- monly with ridgelike to pustulose monticules. Intrazoarial over- growths generally covering large areas of growth surface proximally, commonly becoming less common and more localized distally. Endo- zones are thin and directly adjacent to basal layers or not developed. In exozone, zooecial walls laminate. Laminae diverging orally at high angle from zooecial boundary zone, arching convex orally across cortex and recurving aborally. Zooecial walls having narrowly integrate appearance in cross section. Terminal diaphragms and intermediate diaphragms occurring; basal diaphragms common. Intrazooecial spines sometimes seen. Taxa included. — Only the type species is included here. The internal characters of the second species, R. mamillosa d’Orbigny, assigned to Reptonodicava by d’Orbigny, and species assigned to Reptonodicava by other authors are unknown to me. Discussion. — Bassler (1935) designated the type species of Reptonodicava without comment. In 1953, Bassler placed Reptono- 140 BULLETIN 291 dicava in synonymy with Ceriopora following Gregory (1896) and Canu and Bassler (1920). Comparison of thin-sections of the type species of both genera reveals significant differences in morphology. The zooecial wall of R. globosa is regular and highly symmetrical but relatively irregular and much less symmetrical in C. micropora Gold- fuss, the type species of Ceriopora. R. globosa is characterized by numerous, closely spaced basal diaphragms, but basal diaphragms have not been identified in C. micropora. Intrazoarial overgrowth appears to be a more important means of zoarial increase in C. micropora than in R. globosa. In C. micropora, the exozonal zooecial walls apparently are composed of orally convex Jaminae; in R. glo- bosa, the walls are probably laminate, but the laminae are recurved from the boundary zone. On the basis of these observable differences in morphology, Reptonodicava and Ceriopora are retained here as separate genera. Remarks on wall structure. — Microstructure of the zooecial walls was poorly preserved in all specimens available for study. Often, secondary changes have obscured the boundary between the zooecial wall and secondary infilling of the zooecial chamber. Com- monly, zooecia appear to have granular or sometimes vaguely laminated structure. Well-defined lineations were observed in a few instances. The lineations, initiated at the boundary zone, are broadly arched and orally convex across the cortex. These lineations are interpreted as remnants of primary lamination. The lamination has been observed to occur only in outer thick-walled portions of the zooecial wall. Presumably, it grades proximally with granular tissue, but its true extent is unknown. This wall structure is like that seen in Diplocava; also, growth habits show some similarity. Diplocava, however, has numerous peristomial diaphragms and almost no basal diaphragms. Reptonodicava globosa (Michelin), 1846 Pl. 41, figs. la-h; Pl. 42, figs. 1a-g; Pl. 43, figs. 1, 2, 3a-d; Pl. 44, figs. 1, 2a-c 1821. Non Millepora conifera Lamouroux, Exposition Méthodique des Genres de l’Ordre des Polypiers, des Zoophytes d’Ellis et Solander, p. 87, pl. 83, figs. 6, 7. 1824. Non Millepora conifera Lamouroux, Defrance, Dictionnaire de Science Naturelles, vol. 31, p. 84. 1846. Ceriopora globosa Michelin, Iconographie Zoophytologique, Description par Localités et Terrains des Polypiers Fossiles de France et Pays En- vironnants, p. 246, pl. 57, fig. 5. CerioporIp CycLosToMEs (Bryozoa): NYE 141 1854. Reptonodicava globosa (Michelin), d’Orbigny, Terrain Crétacé Bryo- zoaires, Paléontologie Francaise Description des Animaux Invertébrés, vol. 5, p. 1014. 1896. Ceriopora globosa Michelin, Gregory, Catalogue of Fossil Bryozoa in the Department of Geology, British Museum (Natural History), The Jurassic Bryozoa, pp. 195-7, text fig. 18, p. 196. 1920. Ceriopora globosa Michelin, Canu, and Bassler, U.S. Nat. Mus., Bull. 106, text-figs. 220A-E, p. 678. Type. — Sherborn (1940) cited the repository for the specimens described and illustrated by Michelin in the [conographie Zoologique as the Caen Museum, Caen (Calvados), France. Unfortunately, all of the pre-World War II collections housed at Caen are believed to have been destroyed in the bombardment during the invasion of Normandy. A few of Michelin’s specimens have been preserved at the Muséum National d’Histoire Naturelle, Jardin des Plantes, Paris, but Michelin’s specimens of Ceriopora globosa are not known to be among them (pers. comm. E. Buge, 1969). Michelin did not desig- nate a holotype, and no specimen is known by me to have been designated as the lectotype. Type locality and horizon. — Lebissey, Luc, Ranville (Cal- vados), France; Middle Jurassic, Bathonien. Material studied. — Specimens from D’Orbigny’s collection, identified by D’Orbigny as Reptonodicava globosa, were borrowed from the Muséum National d’Histoire Naturelle, Paris. These speci- mens were almost certainly studied by D’Orbigny for his concept of the genus Reptonodicava, and it is possible that he had com- pared these specimens directly with Michelin’s primary types of Ceriopora globosa. The specimens (d’Orb. Coll. 2988-1 through 5) display the characters described by Michelin, and resemble his illus- trations closely in zoarial shape, surface topography and external appearance of zooecial openings (compare PI. 42, fig. la to pl. 57, figs. 5a, b of Michelin, 1846). In addition, D’Orbigny’s specimens were collected at Luc (Calvados), France, one of three localities cited by Michelin. For the above reasons, D’Orbigny’s specimens are considered to be conspecific with R. globosa in spite of the present impossibility of direct reference to Michelin’s primary types. Thin-sections and acetate peels were made of specimens MNHN d’Orb. Coll. 2988-1 to -3. The card to which the specimens were attached gave the following: Bathonien, Luc (Calvados). Thin- 142 BULLETIN 291 sections and acetate peels were also made from USNM 32171-1 to -5 from the Bathonien, Ranville (Calvados), and USNM 32180-1 to -4, Bathonien, Langrun. Duplicate acetate peels are preserved in the National Museum of Natural History collection and the author’s collection. Description. — Mode of growth — Zoaria relatively large (see Table 25). Basal layers of locally occurring intrazoarial overgrowths are relatively thick (.01 to .02 mm). Exozone — Zooecia often grow continuously for long distances (PI. 43, fig. 3a; Pl. 44, figs. 1, 2a-c). Zooecial walls are symmetrically thickened across zooecial boundary zones. Two major variations are seen locally in the longitudinal profile of the zooecial walls. In one type, zooecial walls show slight longitudinal variation in thick- ness mainly associated with widely flared interzooidal pores, and resulting in moniliform profiles. Monili are commonly circular, oblong or elliptical (PI. 43, fig. 3c), less commonly clavate, obovate or sagittate. The monili show little variation in thickness longi- tudinally resulting in a zooecial chamber with relatively smooth sides overall. When interzooidal pores are uncommon, the zooecial walls locally are parallel-sided (pl. 41, fig. le). On a larger scale, zoaria show repetition of growth zones in which zooecial walls gradually increase in thickness (Pl. 44, figs. 1, 2a-c), producing elongate club-shaped profiles (as modified by local variations described above). Each thick-walled terminal phase is followed by a renewal of thin-walled growth. The boundary zones of major growth phases are commonly marked by concentrations of opaque granules within the wall (PI. 44, figs. 1, 2a-c). Subzones within each major growth phase are marked by thinner concentra- tions of opaque granules in the wall (Pl. 44, fig. 2c). Zooecial chambers commonly have elliptical to subelliptical and smoothly rounded cross sections. Less commonly, short, blunt, intra- zooecial spines extend into the chamber producing crenulate out- lines. Interzooidal pores are numerous and have large diameters (Table 25, ZdPr-MnDr). Diaphragms — Terminal diaphragms occur infrequently and are generally seen subjacent to intrazoarial overgrowths, usually with 143 Cerroporip CycLostomes (Bryozoa): NYE “UOTJOIS jenuasdsur} WO1f IPeW SJUIWIINSEIW $9U0ZOXd TIINO +: [ IUOZOXY SII}JIWIT| [IU UT, § $ SZ Lz £00" Z10" 900° 610" IQuUy-1gpz + £ 99 99 0 y USS9Z /UD-1dPZ z £ SL SL 9b STO" £0" 900° 80° Y.L-LAA2ZPO UX N-4S89-YO2Z z £ SL SL +1 g ZT OT 4 UqXIN-USSD-YO2Z z £ SL SL 61 £0" Be £0" £2" UGX N-USSO-YO2Z UGXIW-USSD-YO2Z z £ SL SL LI +0" 1: 60° 1" ] auozoxg - [2199007 I 6 6 08 061 WW M-IZ [e11e0Z7 apoj eds = IZN -OZN N Act *S «X «WO yayoBIeYyO (NITAHOIN) PSO*OTD FAPIOIGONOLddY AO SLNAWAYXNSVAW AO AYVWWNAS TVOLLSILV.LS $c WI€VL BULLETIN 291 144 "€ ‘@ ‘T-I81ZE WNSM ‘€ ‘Z ‘I-IZIZE WNSQ Sueuads paryruapt {¢ ‘Z ‘I-886¢ NHNW :Ausiqio,q Aq palynuap! suaunsedg "€ ‘I-ILIZE WNSO ‘6 ‘2% ‘I-8862 NHNW "(9T) €- “(S2) Z- ‘(S%) 1-8862 NHNW ‘2-8862 NHNW *(yoea 67) € ‘2 ‘1-886 NHNW aw tw ad0o0 NAWIOddS OL AIM “UON}NIS ASIZASUBI} WOIJ PEW S}UdWIINSvIUI {aUOZOXa IaUU] : 7 aU0ZOXY SId}JIWINT[[IW UT £ T 02 02 0 Ss899Z/4D-1d PZ £ T 02 02 8E 900° $T0° 900° $20° WL-TIM2ZPO IQXWN-USS)-4D9Z £ T 02 02 8 Ee ct hae Sur UdXW-4Ss)-4O2Z £ T 02 02 IT T0° 80° 90° or UGX N-FS8D-4D9Z £ T 02 0d IT 10° or L0° cr UQXIN-4S8)-4YD9Z Z au0zoxy - [8109007 2poj99dg = IZN (OZN N “AO #S aX * WO JoPBIBYO (NITHHOIN) FSOPOTD FAFIIGONOLdAY AO SLNANAYASVAW JO AUVWWAS ‘IVOLLSLLV.LS CERIOPORID CycCLosToMEs (Bryozoa): NYE 145 a small space between the diaphragm and the suprajacent basal layer (Pl. 43, fig. 3d). The diaphragms are commonly planar, about .01 to .02 mm thick, and have a short, aborally curved abutment. Basal diaphragms are thin (about .001 to .002 mm), numerous, and often evenly spaced (PI. 43, fig. 3a; Pl. 44, fig. 2c). The dia- phragms may be slightly arched aborally, laminate with two or three laminae, and arched orally at the juncture with the zooecial wall which forms an abutment distinct from the zooecial wall (Pl. 43, fig. 1). Intermediate diaphragms occur rarely and are seen in the proxi- mal parts of zoaria. The diaphragms are thin (about .001 mm) and arched orally. They flex aborally at the juncture with the wall to merge with the zooecial lining. TABLE 26 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN REPTONODICAVA GLOBOSA (MICHELIN) Cire. __ Ellip. Ov. Pyr. Polyg. Triang. Irreg. Outer Exozone Regular 5 32 Sub 6 17 6 Irregular 1 15 Inner Exozone Regular 10 Sub 4 1 Irregular 5 Discussion. — Michelin described Ceriopora globosa as a new species but cited Muillepora conifera Defrance (1824, p. 84) as a synonym. This reference was investigated because Ceriopora globosa might be considered as a junior subjective synonym of Millepora contfera. Defrance (1824, p. 84) had not erected M. conifera and clearly referred the name to Lamouroux (1821). Defrance included a re- statement of Lamouroux’ original description with the addition of his own observations, presumably drawn from specimens which he identified as M. conifera Lamouroux. Perhaps Michelin was referring to specimens which Defrance described as globular varieties of M. conifera; however, Michelin 146 BULLETIN 291 751A 75 B ~~ ~ (6) oO Ss (= oO feb) %50 250 (oy to Fe is 2 nN on ie) On mm 4a <2 “S) 1.0 2.0 ZcCh-CsSn-NMxDn ZcCh-CsSn-MxDn ZcCh-CsSn-NMxDn 751 a 100 Cc o ron rT) of > - 3250 25 Bi =>! = 25 =e) oO mm 5 | ar 11325.3. 4a 5D no CdZcWI-Th ZdPr-Cn/ZcCsSn Text-figure 17 A-D. Histograms and cumulative curve from three topo- types of Reptonodicava globosa (Michelin). A. Normal to maximum cross- sectional dimension of a zooecial chamber. B. Ratio of the maximum cross- sectional dimension of a zooecial chamber to the normal to maximum cross- sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness D. Count of interzooidal pores per zooecial cross section. CERIopoRID CycLosToMEs (Bryozoa): Nye 147 made no statement to that effect. Therefore, only the specimens described by Lamouroux bear on the possibility of synonymy. Lamouroux’ specimens are presumed lost, and a direct com- parison cannot be made. In this case, however, Lamouroux’ figures are considered to be adequate for comparative purposes. Both the zoarial form and the external appearance of the zooecial openings in M. conifera Lamouroux are different from the same characters as illustrated by Michelin for Ceriopora globosa. Thus, Michelin is considered to have made an error in placing Millepora comtfera Defrance in synonymy with Ceriopora globosa. Genus TETROCYCLOECIA Canu, 1919 Type species: Tetrocycloecia dichotoma Canu, 1919, by original designation, for specimens misidentified by Reuss (1848) as Hetero- pora dichotoma Goldfuss (1826). 1848. Pars Heteropora Blainville, Reuss, Naturwiss. Abh., vol. 2, p. 35. 1919. Tetrocycloecia Canu, Soc. Géol. France, Bull., ser. 4, vol. 17, p. 346. 1920. Tretocycloecia Canu and Bassler, U.S. Nat. Mus., Bull. 106, p. 826. Obj. 1953. recitiele cits Canu and Bassler, Bassler, Treatise on Invertebrate Pale- ontology, Part G, Bryozoa, p. G70. Obj. syn. 1957. Tretocycloecia Canu and Bassler, Buge, Mus. Nat. d’Hist. Nat., Mém., ser. C, Sciences de la Terre, vol. 6, p. 127. Obj. syn. Tentative diagnosis. —Zoaria branching. Branches have exo- zones coaxial with endozones, but intergrading through broad zones of zooecial flexure. Zooecia dimorphic. In outer exozone, zooecial walls indistinctly laminate. Laminae broadly arched, convex orally, and continuous across Zooecial boundary zone. Zooecial lining present. Intrazooecial structures not observed. Taxa included. — Only the type species, Tetrocycloecia dicho- toma Canu, 1919a. The internal characters of other species as- signed to Tetrocycloecia are unknown to me. Discussion. — Canu and Bassler (1920, p. 826) emended Canu’s original spelling of Tetrocycloecia to Tretocycloecia. This emenda- tion has been followed by later authors; Buge (1957, p. 127) be- lieved that the emendation was justified because “Tetrocycloecta résultant d’une erreur de transcription du mot grec ayant servi de base au préfixe du nom du genre.” Canu, however, used the prefix tetro six times in the original publication: 148 BULLETIN 291 a) p. 346, original spelling of Tetrocycloeciadae, a new family, repeated twice. b) p. 346, original spelling of Tetrocycloecia, a new genus repeated in the explanation of pl. 10. c) p. 346, original spelling of Partetrocycloecia, a new genus. Thus, there is no evidence in the original publication that could be interpreted as an inadvertent error by Article 32a, ii, of the ICZN, which specifically excludes incorrect transliteration from considera- tion as an inadvertent error. Evidence for an inadvertent error was not given by Canu and Bassler (1920) when the spelling was emended. The original spelling, Tetrocycloecia, is herein retained; and Tretocycloecia Canu and Bassler (1920) and subsequent authors is considered as an unjustified emendation; therefore, a junior ob- jective synonym of Tetrocycloecia Canu (1919). Tetrocycloecia dichotoma Canu, 1919 Pl. 45, figs. la-g; Pl. 46, figs. la-d, 2a-b, 3 1826. Non Ceriopora dichotoma Goldfuss, Petrefacta Germaniae, vol. 1, p. 34, pl. 10, figs. 9a-e. 1848. Heteropora dichotoma (Goldfuss), Reuss, Naturwiss. Abh., vol. 2, p. 35, pl. 5, figs. 20a, b. Mis-I.D. 1877. Heteropora dichotoma (Goldfuss), Manzoni, I Briozoi Fossili del Mio- cene d’Austria ed Ungheria, Part 3, p. 19, pl. 12, fig. 46. Mis-I.D. 1919. Tetrocycloecia dichotoma Canu, Soc. Géol. France, Bull., ser. 4, vol. 17, p. 346, but mon pl. 10, fig. 10. 1920. Non Tretocycloecia dichotoma Canu, and Bassler, U.S. Nat. Mus., Bull. 106, p. 826, text-fig. 275A-I. Mis-I.D., Inv. Emend. Sp. 1920. ? Tretocycloecia dichotoma Canu and Bassler, Canu, Soc. Géol. France, Bull., ser. 4, vol. 19, p. 213. Inv. Emend. Sp., Mis-I.D.?, no text or plate. 1922. Non Tretocycloecia dichotoma Canu and Bassler, Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 61, pp. 108-10, text-fig. 31A, B. Inv. Emend. Sp., Mis-I.D. 1934. Non Tretocycloecia dichotoma Canu and Bassler, Canu, and Lecointre, Soc. Géol. France, Mém., vol. 9, Part 4, pp. 197-8, pl. 38, figs. 1-14. Inv. Emend. Sp., Mis-I.D. 1957. Non Tretocycloecia dichotoma Canu and Bassler, Buge, Mus. Nat. d’Histoire Naturelle, Mém., ser. C, Sciences de la Terre, vol. 6, pp. 127-9. Inv. Emend. Sp., Mis-I.D. Type. — Six specimens (NMW 1859 L686-1, 2, 3, and NMW 1867xL-1, 2, 3), identified by Reuss as Heteropora dichotoma Gold- fuss, are preserved at the Naturhistorisches Museum, Wien. Specimen NMW 1859 L686-1 bears a close resemblance to Reuss’ figure, 1848, pl. 5, fig. 20 (PI. 45, fig. la). The remaining specimens are consistent in external appearance with Reuss’ descrip- CERIOPORID CycLosToMEs (Bryozoa): NYE 149 tion (1848, p. 35) and figure, and were collected at Eisenstadt, Austria, the type locality. These specimens are not, however, con- sidered here to be syntypic. Reuss commonly listed the date of publication in which a specimen was described, illustrated, or both, as part of the identification number of the specimen (fide A. H. Cheetham). The identification numbers with the specimens studied here do not correspond to any publication of the same date and are considerably later than the original publication in 1848. Although the specimens are not known to be primary types, they are con- sidered to be the most authoritative specimens available. Specimen NMW 1867xL-1 was figured by Manzoni (1877, pl. 12, fig. 46). Type locality and horizon. — Miocene, Tortonian, Leithakalke; Eisenstadt, Austria. Material studied. — The specimens, NMW 1859 L686-1, 2, 3, and NMW 1867xL-1, 2, 3, were kindly loaned by Dr. Heinz Koll- man, Naturhistorisches Museum, Wien. Three specimens were thin- sectioned and peeled: 1859 L686-1. 3 thin-sections, 3 acetate peels. Part of the speci- men remained after sectioning. 1859 L686-2. 3 thin-sections, 3 acetate peels. Part of the speci- men remained after sectioning. 1867xL1-1. 1 thin-section and 2 acetate peels. Duplicate peels are preserved in the National Museum of Natural History collection and the author’s collection. Description. — Mode of growth — Branches are subcylindrical. Zooecia com- monly intersect the zoarial surface at 60° to 80° (PI. 45, fig. 1f; PI. 46, figs. 1b, 2b). Endozone — Zooecia are undulatory in growth. Zooecial walls are generally symmetrically to subsymmetrically thickened across zooecial boundary zones, and are parallel-sided to variably thick- ened with submoniliform to moniliform cross-sections (Pl. 45, fig. lg; Pl. 46, fig. 2b). Zooecial chambers are commonly subelliptical in cross-section. Interzooidal pores are rare, but are large in diameter (about .01 mm). Zooecial walls are homogeneous to subgranular in appearance with a thin zooecial lining. 150 BULLETIN 291 Exozone — Large dimorphs are slightly undulatory and inter- sect the zoarial surface at 60° to 80°. The walls of large dimorphs have apertural rims which project slightly above the zoarial sur- face. Large dimorphs are generally separated by small dimorphs, and commonly are evenly distributed (Pl. 45, figs. la-c, e; Pl. 46, figs. la, c, 2a, 3). Chambers of large dimorphs have elliptical to subcircular cross sections. Zooecial walls in the inner exozone com- monly have subgranular cortices with thin zooecial linings. Lamina- tion becomes more distinct orally, sometimes forming thin zones of laminate tissue which are broadly arched convex orally across the boundary zone. Also, the zooecial lining thickens in the outer exo- zone to about .02 mm. TABLE 28 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN TETROCYCLOECIA DICHOTOMA CANU* Circ! Ellip. Ov. Pyr. Polyg. Triang. Irreg. Large Polymorphs - Exozone Regular 1 3 Sub 1 Irregular Smal] Polymorphs - Exozone Regular 6 35 16 1 Sub 22 10 12 Irregular ce Endozone Regular 7 Sub 3 10 2 2 Irregular 1 *Outlines of exozonal zooecial voids were estimated from NMW 1859 L686-1 (50 zooecia) and NMW 1867xL1-1 (49 zooecia). Outlines of endozonal zooecial voids were estimated from NMW 1859 L686-1. Discussion. — Canu (1919) and subsequent authors mistakenly cited the type species of Tetrocycloecia as Heteropora dichotoma Reuss, 1848. Reuss had identified a suite of specimens from Eisen- stadt as Heteropora dichotoma (Goldfuss), 1826. Canu (1919, p. 346) stated that “Reuss a confondu cette espéce avec Ceriopora dichotoma Goldfuss 1826”, and designated Heteropora dichotoma 151 Nye Cer1oporID CycLosToMEs ( Bryozoa): S1a}9UIT|[TU UT 9 Z c6 26 $c To" $0° 10° 60° UOXIN-U$8D-q09Z auozoxy - sydiow jog [[BuWsg - [e199007 $ c $ $ 0 4 498997 /4O-Fd PZ $ c $ $ Tt Tl0° 9£0° cc0° $s0° YL-TLM9ZPO UGXININSUSFOSNO2Z $ c $ $ 6 or TT OT eT BGAINEUS§D 1992 ] c $ $ 8 TO cr Ir 1 BGXININGUSS OBO 9Z $ c $ $ 9 T0° tr cl +7 UQXIN-YS8)D-YO9Z au0zOxyY - sydiowAlog asieyq - [e109007 + c st st Ot £00° £00° £00° £10" TqQUN=*aPZ 4 c 66 66 0 0% US8D9Z/ Ud PZ c c 66 66 Tf c10" 10° 610° 8Z0° YL-TMPZPO UGXIWIN-USS0-4O9Z c c 66 66 ET c ct at LT BGXINGS§D192Z c c 66 66 Ob 20° $0° 10° a UGX NIN-YS80-YO9Z c c 66 66 8é 20° 90° 10° tT UdXIN-US8)-YOPZ auozoxy - sydiow jog [[V - [B199007 L c 4 61 07 ACG 6.9 Net TiO Dat [Bl1B0Z apoD ‘dads TZN 9ZN N ‘A'O «S *X TIPELEUS ONVO PWOLOHDIC VIDZOTIXIONLAL AO SLNAWAYNSVAW AO AUVWINAS TVOILSILV.LS Le ATaVL "(L¥) T-1TXL98T MWN : (St) 1-989T 6581 MIN °(Z) I-TTXL98T AWN : (£€) 1-989T 6581 MINN *(Z) I-TTXZ98T MN * (8) 1-9897 6881 MIAN *(6%) I-TTXL98T AAINN ‘(0$) 1-989T 6581 MINN ‘I-TIXL981 MIAN ‘Z ‘T-989T 6581 AAWN : (¢2dAjudg) ssnay Aq parsnuep! susumoeds addjodo 7, adqd00 NAWI0ddS OL ATH ‘Z-989T 6S8T *1-989T 6581 MIN ° T-989T 6S8— AAINN aA FHSN SIIJIWN| [IW UTy £ I £ £ 900° 800 IQu\-1gPZ £ I 4 SZ 0 4 USs99Z/4D-1dPZ =~ € T SZ SZ 97 +00" 9T0° O10 9720 UL-LMA2ZPO = UOXxWN-FS807409Z = £ T SZ SZ LI c oT OT V2 UQXIW-USsSD-YDIZ 4 £ i SZ Sc O£ Z0° 80° Z0° II UQXWN-USSD-YDIZ a £ I SZ Sc Te £0° or £0° LT sqoronade sducutana ec aoe 9 Z 26 c6 0 4 USSD2Z/UD-1dPZ 9 4 26 26 O€ £10° T+0° 610 840 U.L-LAM2ZPO UQXWN-4S80-4O2Z 9 cz C6 26 cl ST ot OT LT UQXWN-4Ss8)-YOIZ 9 c c6 26 9¢ T0° +0° 10° 80 UQXIWN-USSD-YOOZ apon v2dg_ IZN (OZN N “AO «8 aX «WO rayoRIeYD 152 ONVO PWOLOHIDIG FPIDAOTIAIONLAL AO SLNAWAUANASVAW AO AUVNWAS TVOLLSILV.LS CERIOPORID CycLosToMEs (Bryozoa): NYE on N oO oO QD frequency NO Oo mm 4a oo) 3 ZcCh-CsSn-NMxDn oO N [e) on QO frequency NO oy) mm I JZ CdZcWI-Th ~ ao jee) Oo ie) uency freq 25 1.0 2.0 ZcCh-CsSn-Mx Dn ZcCh-CsSn-NMxDn oO N ©) oO ie) oO cumulative % 12534) 50 ZdPr-Cn/ZcCsSn 153 Text-figure 18 A-D. Histograms and cumulative curve from three topo- types of Tetrocycloecia dichotoma Canu. A. Normal to maximum cross-sectional dimension of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Compound zooecial wall thickness. D. Count of interzooidal pores per zooecial cross section. 154 BULLETIN 291 Reuss as the type species of Tetrocycloecia. By Article 70B of the ICZN, the primary type specimens of Tetrocycloecia dichotoma were the specimens before Reuss, but by Article 70B, the name assigned to those specimens is Tetrocycloecia dichotoma Canu, 1919. Gregory (1909, p. 199, footnote 2) believed that Ceriopora dichotoma Goldfuss and T. dichotoma Canu belonged to different genera. This opinion was based on external characters seen in speci- mens of C’. dichotoma Goldfuss from the type locality. Canu and Bassler were of the same opinion, and in 1922 (pp. 119-20, text- fig. 35) described and illustrated specimens identified as conspecific with C. dichotoma Goldfuss from the type locality for which they erected the genus Grammascosoecia. The secondary specimens differ internally in many characters, ¢.g., bifoliate growth habit, from T. dichotoma Canu as understood here. Until the internal characters of the primary types of C. dichotoma Goldfuss have been studied, there must remain some question as to the reliability of existing species concepts as applied to that name. The species concepts, as presently understood, and the temporal separation between C. dichotoma Goldfuss and T. dichotoma, however, provide reasonable grounds for continuing to consider C. dichotoma and T. dichotoma Canu as separate entities. Canu and Bassler, 1922, p. 108, noted that: “. . . the studies relative to this species have been made from specimens collected in France. We are not entirely certain of our determinations, for we have never been able to procure Austrian specimens for comparison.” A comparison of Reuss’ specimens of 7’. dichotoma to specimens from the Miocene of France figured and identified by Canu and Bassler and later authors, reveals a number of morphological differences (see Canu and Bassler, 1920, text fig. 275A-I; Canu and Bassler, 1922, pp. 108-110, text figs. 31A, B; Canu and Lecointre, 1934, pp. 197-8, pl. 38, figs. 1-14; Buge, 1957, pp. 127-9). The following charac- ters identified in the non-Reuss specimens are not present in the Reuss specimens: 1) Well-defined boundary between exozone and endozone. 2) Zooecia intersect surface at approximately 90°. 3) Endozone walls nonmoniliform. CERIOPORID CycLosToMEs (Bryozoa): NyY& 155 4) Basal diaphragms in endozonal zooecia. 5) Intermediate diaphragms in exozonal zooecia. 6) Zooecial walls in exozone subsymmetrical to asymmetrical in thickness across the boundary zone. The specimens from the Miocene of France described by Canu and Bassler (1920, 1922), Canu and Lecointre (1934), and Buge (1957), are here not considered to be conspecific or congeneric with T. dichotoma Canu because of the morphotypic differences described above. Genus ZONOPORA d’Orbigny, 1849 Type species: Ceriopora spiralis Goldfuss, 1826, p. 36, by orig- inal designation and monotypy, D’Orbigny (1849, p. 503). 1849. Zonopora d’Orbigny, Rev. et Mag. Zoologie, vol. 1, ser. 2, p. 503. 1854. Spiroclausa d’Orbigny, Terrain Crétacé Bryozoaires, Paléontologie Fran- caise Description des Animaux Invertébrés, vol. 5, p. 883. Obj. syn. 1909. Zonopora d’Orbigny, Gregory, Catalogue of Fossil Bryozoa in the De- partment of Geology, British Museum (Natural History), The Cretaceous Bryozoa, vol. 2, p. 427. 1922. Spiroclausa d’Orbigny, Canu, and Bassler, U.S. Nat. Mus., Proc., vol. 61, p. 92. Obj. syn. 1953. Zonopora d’Orbigny, Bassler, Treatise on Invertebrate Paleontology, Part G, Bryozoa, p. G71. Tentative diagnosis.—Zoaria branching; zooecia dimorphic. Large zooecia bud sequentially in helical pattern. Large and small zooecial apertures arrayed in parallel, helical zones. Zone of large zooecial apertures forming a continuous zoarial salient; zone of small zooecial apertures forming a continuous zooecial embayment. Local intrazoarial overgrowths occur occasionally. In exozone, outer cortex of zooecial wall light-colored and granular to indistinctly laminate. Laminae broadly curved, convex orally, but commonly not continuous across boundary zone. Zooecial lining thick, dark-colored, with longitudinally directed, undulatory, sometimes crenulate laminae. Terminal diaphragms and mural spines occurring. Taxa included. — Only the type species. Internal characters of the types of other species commonly assigned to Zonopora are un- known to me. 156 BULLETIN 291 Zonopora spiralis (Goldfuss), 1826 Pl. 47, figs. la-g; Pl. 48, figs. la-g; Pl. 49, figs. la-e; Pl. 50, figs. la, b, 2a-c, 3 1826. Ceriopora spiralis Goldfuss, Petrefacta Germaniae, vol. 1, p. 36, pl. 11, 1849. Bees eaeoira (Goldfuss), d’Orbigny, Rev. et Mag. Zoologie, vol. 1, 1850. Dae i spiralis (Goldfuss), d’Orbigny, Prodrome de Paléontologie Stratigraphique Universelle, vol. 2, p. 267. 1854. Spiroclausa spiralis (Goldfuss), d’Orbigny, Terrain Crétacé Bryozoaires, Paléontologie Francaise Description des Animaux Invertébrés, vol. 5, p. 883, pl. 764, figs. 1-5. 1899. Zonopora spiralis (Goldfuss), Gregory, Catalogue of Fossi] Bryozoa in Department of Geology, British Museum (Natural History), The Cre- taceous Bryozoa, vol. 1, pp. 427-8. 1922. Spiroclausa spiralis (Goldfuss), Canu, and Bassler, U. S. Nat. Mus., Proc., vol. 61, p. 92-4. Type. — UB 133 is here designated as the lectotype. The lecto- type, labeled “Original zu Goldfuss, 133, Ceriopora spiralis”, was firmly cemented to a piece of coquinoid limestone primarily com- posed of worn bryozoan fragments, echinoid spines and various shell debris. The lectotype is similar in appearance to the specimen figured by Goldfuss, 1826, pl. XI, figs. 2a, b. Type locality and horizon. —“Petersberge bei Maastricht;” rocks presently exposed here are Upper Cretaceous, Maastrichtian in age. Material studied. — Dr. A. Durkoop, Geologische Palaeonto- logische Institut of Bonn, kindly made available the lectotype speci- men UB 133. Three thin-sections and four acetate peels on a single acetate slide were made from the lectotype; approximately one-half of the lectotype remained after sectioning. Four topotypes, collected in the Upper Maastrichtian from Quarry Curfs near Maastricht, were kindly made available for study by Prof. E. Voigt. In addition, 15 topotypes in the collection of the National Museum of Natural History were thin-sectioned and peeled. The specimens were labeled as “Cretaceous, Maastrichtian, Maastricht, Holland, USNM Loc. 2965A”. Duplicate acetate peels of all specimens sectioned are in the collections of the National Museum of Natural History and the author’s collection. Description. — Mode of growth — Branches have subcircular to elliptical cross sections. Distal to branch bifurcations, the helix pattern of budding Certoporip CycLosToMEs (Bryozoa): NYE 157 is reversed in one branch. New branches sometimes arise as over- growths (PI. 48, fig. 1a). Endozone — Zooecial walls are thin and parallel-sided. Zooecial chambers commonly have subpolygonal or, less commonly, subellipti- cal cross sections. Interzooidal pores are rarely seen. Zooecial walls are light-colored and homogeneous to subgranular with thin zooecial linings. Exozone — Zooecial walls are symmetrical in thickness, or near- ly so, across the zooecial boundary zone. Zooecial walls are sometimes nearly parallel-sided (PI. 50, figs. la, b, 3) but more commonly show regular increase in thickness to a maximum which is located slightly suboral to the aperture, producing lancet (PI. 48, figs. Ic, d, e) or clavate (Pl. 50, fig. 2a) profiles. Locally, most commonly in inner exozone portions of the wall, submonoliform (PI. 48, fig. le; Pl. 49, fig. le) and sometimes moniliform cross sections are seen. These result from relatively slight thinning of the wall close to interzooidal pores. Interzooidal pores are narrow (about .01 mm in diameter), straight, and cylindrical for most of their length (PI. 48, fig. 1g; Pl. 49, fig. le). Zooecial chambers commonly have elliptical to sub- elliptical cross sections. Mural spines are narrow (about .006 mm in diameter), locally numerous (PI. 48, fig. 1d; Pl. 49, fig. le; Pl. 50, fig. 3), and are most commonly seen in the exozone. The spines have a light-colored core continuous from light-colored tissue in the zooecial wall (PI. 50, fig. 3). In older zooecia, the zooecial lining commonly overlaps and buries the spines (PI. 49, fig. le). Diaphragms — Terminal diaphragms are numerous and com- monly occur in smaller zooecia. The diaphragms are thin (about .02 mm), and pores are about .01 mm in diameter. The walls of zooecia with terminal diaphragms often have constricted apertural rims (PI. 50, figs. la, 2a, b, c). The calcareous tissue of the diaphragms (except where interrupted by pseudopores) is structurally continuous with the zooecial lining of the apertural wall (Pl. 50, figs. 2b, 2c). The oral surfaces of the diaphragms are flush with the apertures and often appear to form sheetlike deposits when viewed externally (PI. 47, figs. la, b). BULLETIN 291 158 "VS96Z ‘907 INS) ‘sedAjodo} g puke get qa edAjoyoaT *Z NAWIOddS OL ATH "EST GQ edAj0VI7T ‘Tf S1OVOUIN| [TUE UT y T T $c $d 9c $00° +00" c00° Lé0° YL-FTIM9Z I I $c $c tb 800° 810° O10" $£0° YL-TM9ZPO T I Sc $c 0 al! USs99Z/UO-1dPZ UQXWN-YSS80-YO9Z I I $c $c El 9 eT OT sh! BGXIN-FS§O S097 I I $c $c tT c0° ae 80° st UGX N-YSS80-Y4O9Z I T $c $c LT Z0° st OT 0c UGXIN-USs0- YZ auozopuy - [8199007 T it 9¢ 9 0 Y USsD9Z /UD-1d PZ I I 9 9¢ tr Z£0 cL0 $20" cot YL-LM9ZPO UGXWN-YS8S0-4D9Z T T 9¢ 9 x4 se st OT £7 UGXW-USs8)-YO2Z I I 9¢ 9 Lb $0 Or £0° Lv UGXINN-YS80-YO9Z T T 9€ 9€ tb 90 +1 +0 Lo UQXIN-US§D-YO2Z U0ZOXY - [EIN9007 Z 6 6 02 v 6T vT Le Udx*W-USs8O-1a [el1v0Z epop eds) =IZN =(OZN N PN) #S aX «WO IayeBIeYO (SSQHd1OD) SITFUldS FXOMONOZ AO SLNANAUANSVAW AO AUVWWAS TVOILSILVLS 6c ATAVL CerR1oporIp CycLosToMEs (Bryozoa): NYE 159 TABLE 30 FREQUENCY OF VISUALLY ESTIMATED OUTLINES OF ZOOECIAL CHAMBERS IN THE LECTOTYPE OF ZONOPORA SPIRALIS (GOLDFUSS) Cire. —_ Ellip. Ov. Pyr. Polyg. Triang. Irreg. Exozone Regular 14 3 1 Sub 1 12 3 1 Irregular 1 Endozone Regular 1 1 2 Sub 2 8 1 10 Irregular Remarks on morphology.— The helical growth habit of Z. spiralis poses particular problems of description and interpretation not encountered with other growth habits. One such problem con- cerns measurement and interpretation of zooecial characters. Certain numerical observations of exozonal zooecia, such as the diameters of the zooecial void, are gathered from tangential sections. The as- sumptions generally made in the interpretation of such measurements are: 1) If the section is tangential to the surface, all zooecial inter- sected by the plane of the section will be at approximately the same ontogenetic stage. 2) All zooecia within a given section intersect the section at ap- proximately the same angle (commonly about 90°); further- more, the angle of intersection with the section is about equal to the angle at which zooecia intersect the zoarial surface. Measurements made from tangential sections of Z. spiralis are generally not consistent with either assumption. Measurements, and the statistics generated from those measurements, must be con- sidered biased and only broadly useful in comparison to species satis- fying the above assumptions. The bias arises from the helical shape of the branches (refer to Text-fig. 2D). In Z. spiralis, surficial sec- tions parallel to the major axis of branch growth are tangential to the zoarial surface only at the latter-most surface of a zoarial salient ( AA’ in Text-fig. 20) or, if a deep section is made, are tangential only to the surface at an embayment (BB’ in Text-fig. 20). 160 BULLETIN 291 751A 75 B ~> ~> oO oO (= (= 250 25 lox on v 2 25 25 mm 4d oe se) 1.0 20 ZcCh-CsSn-NMxDn ZcCh-CsSn-MxDn ZcCh-CsSn-NMxDn ah on N (o) fo) uo (o) Nh on cumulative % ese ler 1a) ZdPr-Cn/ZcCsSn Text-figure 19 A-C. Histograms and cumulative curve from the lectotype of Zonopora spiralis (Goldfuss). A. Normal to maximum cross-sectional dimen- sion of a zooecial chamber. B. Ratio of the maximum cross-sectional dimension of a zooecial chamber to the normal to maximum cross-sectional dimension of a zooecial chamber. C. Count of interzooidal pores per zooecial cross section. CeRIoporRID CycLosTomMEs (Bryozoa): NYE 161 f. 35522 I// : ees ZOOCIa // small small @ : boecia eee re Hh, A B ic Text-figure 20 A-C. Budding pattern of Zonopora spiralis (Goldfuss). A. An external view of a branch to show the arrangement of large and small zooecia in parallel, helically coiled zooecia. Small zooecia are commonly covered by terminal diaphragms. B. Idealized longitudinal section. Zooecia are curved only in profile through part of any section. In order to show their arrangement, more of the zooecial length is shown in this figure than would be commonly intersected in an actual section. AA’ shows the position of a shallow tangential section which intersects only large zooecia. BB’ is a deep tangential section — large zooecia in deep tangential section, but small zooecia in shallow tangential section. C. Cut away view of a branch. Branch outline indicated by dashed line. Only a single sequentially budded series of large zooecia is shown. Loci of budding indicated by dark helix in central area of branch. o 162 BULLETIN 291 In addition, sections taken along AA’ intersect only zooecia with large cross sections; sections taken at BB’ intersect thin- walled zooecia with large diameters and clusters of thick-walled zooecia with small diameters. In nearly all sections in which the plane is parallel to the major axis of distal growth, zooecia intersect the plane of the section at a different angle than that at which they intersect the surface. For example, note that zooecia with large cross sections in the middle righthand salient intersect a surface approxi- mately coincidental with CC’, rather than at a plane parallel to the major axis of branch growth. When viewed externally, terminal zooecial coverings sometimes appear to have coalesced as a single calcareous sheet over the zooecial apertures (PI. 47, figs. la, b). If this were so, the calcareous tissue would have been deposited from an outer membrane (gymnocyst of Borg). Thin sections reveal, however, that the diaphragms are typical terminal diaphragms. The diaphragms are not superposed over the zooecial walls but are structurally continuous. The dia- phragms were emplaced by tissues aboral to the diaphragm. REFERENCES CITED Banta, W. C. 1969. The body wall of cheilostome Bryozoa, Part 2. Interzoidal com- munication organs. Jour. Morphology, vol. 129, pp. 149-170, 26 text-figs. Barrois, Jules 1877. Récherches sur ’embryogénie des bryozoaires. Travoux de |’Institut Zoologique de Lille et de la Station Zoologique de Wimereux, vol. 1, pp. 1-305. 1879-1880. Mémoire sur la metamorphose des bryozoaires. 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Michelin, Hardouin 1841-1848. Iconographie Zoophytologique description par localités et ter- rains des Polypiers fossiles de France et Pays Environnants. Pp. 1-348, 79 pls. Dates fide Sherborn in Gregory, 1909, p. 23: p. 1-40, 1841 p. 145-184, 1845 p. 41-72, 1842 p. 185-248, 1846 p. 73-104, 1843 p. 249-328, 1847 p. 105-144, 1844 p. 329-348, 1848 Nicholson, H. A. 1880. On the minute structure of the Recent Heteropora neozelanica Busk, and on the relations of the genus Heteropora to Monticuli- pora. Ann. Mag. Nat. Hist., ser. 5, vol. 6, pp. 329-423, 5 text-figs. 166 BULLETIN 291 Nielsen, Claus 1970. On metamorphosis and ancestrula formation in cyclostomatous bryo- zoans. Ophelia, vol. 7, pp. 217-256, 41 text-figs. Nye, O. B., Jr. 1969. Aspects of microstructure in post-Paleozoic Cyclostomata. In Pro- ceedings of the First International Bryozoology Association Con- ference, Milan, Italy, Soc. Italiana Sci. Nat. Milano, Atti, vol. 108, pp. 111-114. _, Dean, D. A., Hinds, R. W. 1972. 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Illustrations of the geology of Yorkshire or, A description of the strata and organic remains of the Yorkshire Coast. Pp. i-xvi, 1- 192, pls. 1-14, printed for the author by Thomas Wilson and Sons, High Ousegate, York. Reuss, A. E. von 1848. Die fossilen Polyparien des Wiener Tertidrbeckens. Ein mono- graphischer Versuch. Naturwiss. Abh., Bd. 2, pp. 1-109, pls. 11. Robertson, Alice 1903. Embryology and embryonic fission in the genus Crisia. Univ. Cali- fornia, Pub. Zoology, vol. 1, No. 3, pp. 115-156, pls. 12-15, Ph.D. dissertation. 1910a. Bryozoans. Papers from the Harriman Alaska Expedition, the Bryozoa, Part VI, Washington Acad. Sci., Proc., vol. 2, pp. 315-334. 1910b. The cyclostomatous Bryozoa of the West Coast of North America. Univ. Calif., Pub. Zoology, vol. 6, pp. 225-284, pls. 18-25. Sherborn, C. D. 1940. Where is the —————— Collection? Pp. 7-149, Cambridge Univer- sity Press, London. Silén, Lars 1944. On the formation of the interzoidal communications of the Bryozoa. Zoologiska Bidrag fran Uppsala, vol. 22, pp. 433-488, 1 pl., 59 text-figs. Tavener-Smith, Ronald 1969. Skeletal structure and growth in the Fenestellidae (Bryozoa). Paleontology, vol. 12, pp. 281-309, pls. 52-56, 9 text-figs. Ulrich, E. O., and Bassler, R. S. 1907. Bryozoa. In Weller, Stuart, A report on the Cretaceous Paleontology of New Jersey. Geol. Sur. New Jersey, Paleont., ser., vol. 4, pp. 313-355, pls. 20-26. CERIOPORID CycLosToMEs (Bryozoa): NYE 167 Utgaard, John 1968a. A revision of North American genera of ceramoporoid bryozoans (Ectoprocta). Part I, Anolotichniidae. Jour. Paleont., vol. 42, pp. 1033-1041, pls. 129-132. 1968b. A revision of North American genera of ceramoporoid bryozoans (Ectoprocta). Part 2, Crepipora, Ceramoporella, Acanthoceramo- porella, and Ceramophylla. Jour. Paleont., vol. 42, pp. 1444-1455, pls. 181-184. 1969. A revision of North American genera of ceramoporoid bryozoans (Ectoprocta). Part 3. The ceramoporoid genera Ceramopora, Pipillalunaria, Favositella and Haplotrypa. Jour. Paleont., vol. 43, pp. 289-297, pls. 51-54. , and Boardman, R. S. 1965. Heterotrypa Nicholson, 1879, and Peronopora Nicholson, 1881 (Bryozoa, Trepostomata): proposed designation of a type-species in conformity with generally accepted usage. Z. N. (S.) 1693. Bull. Zool. Nomenclature, vol. 22, pp. 112-118, 1 table. Viskova, L. A. 1965. Late Cretaceous Bryozoa of the genus Meliceritites from the Middle Volga region. Paleont. Zhur. No. 3, pp. 49-58, 2 pls. 1968. New bryozoans of the suborder Salpingina (Novyye mshanki podu- tryada Salpingina). Paleont. Zhur., No. 2, pp. 32-41, 4 text-figs. Voigt, Ehrhard 1953. Revision von; H. Hamm, Die Bryozoen des Maastrichter Obersonon (1881). Geol. Staatsinst. Hamburg, Mitt., vol. 22, pp. 32-75, pls. 1-14. 1971. The cheilostomate nature of the alleged cyclostomatous bryozoan genus Dysnoetopora. Lethaia, vol. 4, pp. 79-100, 10 figs. Waters, A. W. 1879. On the occurrence of Recent Heteropora. Royal Micro. Soc., Jour. Vol. 2, pp. 390-393, pl. 15. 1884a. Closure of the cyclostomatous Bryozoa. Jour. Linnean Soc., London, Zoology, Jour., vol. 17, pp. 400-404, pl. 17. 1884b. Fossil cyclostomatous Bryozoa from Australia, Quart. Jour. Geol. Soc., Nov. 1884, pp. 674-697, pls. 30, 31. 1890. On some ovicells of cyclostomatous Bryozoa. Linnean Soc. London, Zoology, Jour., vol. 20, pp. 275-285, pl. 15. = pine. 9&4 sh* “¢ ws th ie oh WIG her a ma te ‘, 7 : : o oan 7 : . i. a> ea cman rent pearriitinesk pee ay path, ha, eee “higw Vadhesiaane mp thom Ly Pniaapry ies, + Pais 92d dt Ba SH i etiOlad Gini! iMesh hee CEL ale tegen amege ahd te ey Ce . ye rips Agaey x mm, Het ation Awe eelske > ki ais Liv areas ¢ é eh Nec *. +1 anes is wh eM Pela fatslo. nay eee!) Cas Billa GA Ara err 4 -! ' OS -Gak - a. f 7 rs bof , dajy* nae ww ar oe one & £ =?) » t WAG Vit : y apes fj ; : Ra | y ie 4 a ; , ‘ ot é ‘iy on 7 an » uy ry » az ~— 1 ’ bar a ; c rf i f H ' ul qo ) ; ' | xi . , ‘ I | “A i ; : 4 ? iV PLATES 170 Figure fe BULLETIN 291 EXPLANATION OF PLATE 1 Ceriocava corymbosa (Lamouroux), 1821 ...............c eee cee MNHN IP2-1, Jurassic, Bathonien; Ranville (Calvados), France. la. 1b. fic: 1d. le. If. lg. ih. Branch 1 with uneven monticular surface; holes are borings. Surface of same branch X10, apparently showing occur- rence of large and small polymorphs. Histogram, however, of cross-sectional dimension of zooecial chambers (text-fig. 6A) has unimodal distribution approaching a normal curve. Surface of same branch X10. Note pattern of zooecial open- ings in upper right monticule and the concentration of smaller zooecia in intermonticular area in center of figure. Longitudinal thin-section of same branch X10. Cyclic growth pattern in endozone visible as center of thin, convex zones, caused by annular thickenings of zooecial walls. Note the very thin, undulatory walls and relatively small zooecial diameters in the endozone. Both zooecial walls and zooecial diameters increase in exozone. Tubular structure along branch axis is a boring. Tangential thin-section of same branch X10, with normally appearing zooecia in top right and bottom left, and zone of irregular budding in central part of figure. Transverse thin-section of same branch X10; all zooecia with terminal diaphragms and thin, closely spaced basal diaphragms. Brood chamber in center of figure; most of endozone removed by a boring organism. Longitudinal thin-section 30. Note circular to oblong moni- liform profiles of zooecial wall in inner exozone, and basal diaphragms becoming more numerous and closely spaced in the exozone. Tangential thin-section 30. Indistinct laminations concen- tric with zooecial chamber. BuLL. AMER. PALEONT., VOL. 69 PLATE 1 BuLL. AMER. PALEONT., VOL. 69 PLATE 2 UL LP Ay Se AO Ee 8 4 pis Pees Deo e casita bani a ALE a Figure il. CERIOPORID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 2 Ceriocava corymbosa (Lamouroux), 1821 0.0.0.0... MNHN IP2-2, Jurassic, Bathonien; Ranville (Calvados), France. las 1b. 1c. 1d. le. tf 1g. 1h. Branch X1. Surface of same branch X5. Surface of same branch X30. Tangential thin-section 10; zooecial walls commonly amalgamate, occasionally integrate. Longitudinal thin-section X10. Thin-walled zooecia with relatively small diameters in endozone. Thick-walled zooecia with large diameter in exozone. Note thick terminal dia- phragms and thin, closely spaced, basal diaphragms in exo- zone. Transverse thin-section 10. Illustrates increase in zooecial diameters from endozone to exozone. Note laminate walls in exozone and terminal diaphragms. Slightly oblique, longitudinal thin-section X10 with brood chamber in lower left portion of figure. Large cavity to right is a boring. Longitudinal thin-section 10. Axillary zone of zooecial intergrowth is shown in upper central part of figure. Zone is just distal to bifurcation of old branch in lower central part of figure. Growth axis of each new branch extends from bot- tom center to left and right hand corners respectively. Much of endozone in right-hand branch cut by mud-filled boring. Note distal flexure and anastomosis of zooecia growing towards each other from each new branch. Cyclic growth in endozone is illustrated in left-hand branch. Each cycle con- sists of zooecial walls with long, thin-walled portion capped by annular thick-walled portion; probably represents a single episode of growth. 171 172 Figure BULLETIN 291 EXPLANATION OF PLATE 3 1-3. Ceriocava corymbosa (Lamouroux), 1821 200.000.0000... All specimens figured from Jurassic, Bathonien; Ranville (Cal- vados), France. iL, la. 1b. ile MNHN IP2-1. Longitudinal section, acetate peel 50; zooecial growth direction to the right. Local zone of irregular budding. Zoo- ecial cavities are partitioned by a thick, non-porous dia- phragm, unlike numerous basa] diaphragms. New zooecial walls are continuous with the oral surface of the diaphragm. Zooecial walls have moniliform profiles, occasiona] mural spines, and thin basal diaphragms. Calcareous tissue of most of cortex is dark in color; outer cortex is composed of light- colored tissue outlining zooecial boundary zone. Note numer- ous basal diaphragms. Tangential thin-section from same zoarium showing zone of irregular budding. Longitudinal section, acetate peel 30. Abandoned brood chamber has floor with interzooidal pores and thick porous roof. Roof is overlain by thin, basal layer and thin-walled ZOoecia. MNHN IP2-2. Transverse section, acetate peel X30. Growth direction of zooecia is to the right in zone of irregular bud- ding. Zooecia are thin-walled and irregular in growth direc- tion. Note large open area in central portion of figure. USNM Cat. 68941-2, tangential section, acetate peel 30. Large cavity is interpreted as a brood chamber. Many of the zooecial walls in this figure have an integrate appearance. BULL. AMER. PALEONT., VOL. 69 PLATE 3 BULL. AMER. PALEONT., VOL. 69 PLATE 4 ‘AE BSA 2 ae fs 7% th rg a ke tae ee re ae ee - “net Figure 1p CERIOPORID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 4 Ceriocava corymbosa (Lamouroux), 1821] 0000.......occoccccccceceeeeeeeees MNHN IP2-2, Jurassic, Bathonien; Ranville (Calvados), France. la. 1b. lie; 1d. le. Tangential section, acetate peel X30. Zooecial walls general- ly amalgamate in appearance. Many zooecia have small mural spines. Longitudinal thin-section 50; detail of Pl. 2, fig. 1h, show- ing anastomosis of orally growing zooecial walls in axillary zones just distal to the bifurcation of a branch. Although zooecial chambers pinch out, zooecial walls merge without break and continue to grow orally. Transverse thin-section 50. Laminae generally indistinct, but broad V-shaped patterns with apices pointing towards zoarial surface can be seen. Note subapertural position of thick terminal diaphragms. Longitudinal thin-section 30. Zooecial walls have alate to sagittate monilar profiles and are indistinctly laminate. Laminae form V-shaped patterns pointing toward zoarial surface. Note correspondence of V-shaped pattern to profile of zooecial wall at skeletal aperture. Basal diaphragms closely and regularly spaced. Longitudinal section, acetate peel 30. In endozone, zooecial walls consist of longitudinally repeated two-stage growth cycles: a long, thin-walled growth stage, and a short stage with annular thickenings. 173 174 Figure BULLETIN 291 EXPLANATION OF PLATE 5 1-3. Ceriocava corymbosa (Lamouroux), 1821 ............00ccccceeccccceeeeeeees 1 la. 1b. Ic. MNHN IP2-1, Jurassic, Bathonien; Ranville (Calvados), France. Transverse section, acetate peel 100. Shows thick terminal diaphragms with aborally flexed abutments apparently not merging with zooecial wall. Poorly defined, light-colored patches indicate probable position of pores. Light-colored tissue in outer cortex outlines zooecial boundary zone. Dark- colored tissue, making up most of cortex, is indistinctly laminated. Laminae generally form broad V-shaped patterns with apices pointing towards the zoarial surface. Transverse section, acetate peel 200. Shows three thin, in- termediate diaphragms just aboral to zooecial aperture. Dark lines sloping aborally from zooecial boundary zone probably represent primary laminate structure. Note granular to near- ly structureless calcite along zooecial boundary zone. Longitudinal section, acetate peel 200. Zooecial wall in exozone shows indistinct laminate structure. Note light- colored zone along zooecial boundary zone. Orally flexed basal diaphragm is seen in lower right. USNM 32164-3, Jurassic, Bathonien; St. Aubin (Calvados), France. Longitudinal section, acetate peel X100. Slightly un- dulatory, nearly parallel-sided zooecial walls in endozone. Walls are granular. Basal diaphragms flex orally at junc- - ture with zooecial wall and continue orally as zooecial lining for a considerable distance; but commonly, not extending to next diaphragm. MNHN IP2-2, Jurassic, Bathonien; Ranville (Calvados), France. Tangential section, acetate peel 100. Moniliform profile of zooecial wall shown here results largely from widely flaring profile of interzooidal pores. Section parallels axis of uppermost interzooidal pore, but is slightly oblique or abaxial to other pores which, therefore, appear to be sealed at the zooecial boundary zone. Vaguely spine- shaped masses of light-colored tissue project toward the zooecial chamber from the outer cortex, but show little (lower half of figure) or no surficial relief. PLATE 5 BULL. AMER. PALEONT., VOL. 69 BULL. AMER. PALEONT., VOL. 69 PLATE 6 Figure CeRIoporID CycLosToMEs (Bryozoa): Nye EXPLANATION OF PLATE 6 1-3. Ceriocava corymbosa (Lamouroux), 1821 .00000000...cceeceeeeeee i. 3a. 3b. 3c. USNM 32164-3, Jurassic, Bathonien; St. Aubin (Calvados), France. Longitudinal section, acetate peel 100. Laminae have broad, V-shaped forms pointing towards the zoarial surface. Light-colored obscurely structured tissue in the outer cortex marks the zooecial boundary zone. Walls are nearly parallel-sided, except where indented by interzooidal pores. MNHN IP2-2, Jurassic, Bathonien; Ranville (Calvados), France. Tangential section, acetate peel 100. Zooecial walls show indistinct lamination; thin zooecial linings; short, bluntly rounded zooecial spines; and interzooidal pores with relatively large diameters. MNHN IP2-1, Jurassic, Bathonien; Ranville (Calvados), France. Longitudinal section, acetate peel 200. Thick terminal diaphragm is thick and slightly subapertural in position. Basal diaphragms are, by comparison, much thinner. Zoo- ecial walls are indistinctly laminate with laminae orally ob- lique at a generally higher angle than those in fig. 1. Transverse section, acetate peel X200. The inner cortex is composed of indistinctly wavy to crenulate laminae. Granu- lar to homogeneous calcite in the outer cortex marks the zooecial boundary zone in the inner exozone. Transverse section, acetate peel X200. The large chamber is a brood chamber. Subjacent zooecial walls and brood cham- ber floor are structurally continuous, forming compound wall. The floor is pierced by pores similar in appearance to in- terzooidal pores. The pores in the roof are sealed proxi- mally by a thin layer of dark-colored tissue which appears to line most of the chamber. 175 176 Figure BULLETIN 291 EXPLANATION OF PLATE 7 1. ‘Ceriopora micropora Goldfuss, 1826 ......0...... eee All views are made from the lectotype, UB 119. This specimen was figured by Goldfuss, 1826, pl. 10, figs. 4a, d; von Hagenow, 1851, pl. 5, fig. 13 as Heteropora crassa; and Voight, 1953, pl. 2, fig. 4 as Pennipora beyrichi Hamm. la. 1b. Ic. 1d. le: if. Zoarial fragment 2; holes in surface are borings. Surface of zoarial fragment 30. Note large variation in size and shape of zooecial apertures. Size variation is con- tinuous, and approaches an unimodal normal curve (Table Ne Tangential thin-section 30. Note large variation in size and shape of cross sections of zooecial chambers. Zooecial walls are broadly amalgamate. Transverse thin-section X10 from proximl portion of zoar- ial fragment. Shows main growth and a single intrazoarial overgrowth. Longitudinal thin-section 30; typical appearance of zoo- ecial walls in thin-walled exozone phase. Diaphragm-like structures in oral portions of two zooecia, to right of center, are fractures in the polyester used to imbed the specimen prior to sectioning. Longitudinal thin-section 30. BuLL. AMER. PALEONT., VOL. 69 PLATE 7 PLATE 8 BULL. AMER. PALEONT., VOL. 69 Figure ie CERIOPORID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 8 Ceriopora micropora Goldfuss, 1826 00.0000... ecteeteeteees All views are made from the lectotype, UB 119. This specimen was figured by Goldfuss, 1826, pl. 10, figs. 4a, d; von Hagenow, 1851, pl. 5, fig. 13 as Heteropora crassa; and Voight, 1953, pl. 2, fig. 4 as Pennipora beyrichi Hamm. la 1b. es 1d. Tangential section, acetate peel 100. Large, primary inter- zooecial chamber (brood chamber?). Longitudinal section, acetate peel X100. Thin-walled, exo- zonal growth shows symmetrically thickened, submonili- form to nearly parallel-sided zooecial walls. Transverse section, acetate peel 100. Zooecia grow toward the right side of figure. Thin-walled endozonal zooecia are continuous from thick-walled zooecia; intermediate basal layer is not seen here. Some zooecial cavities are continuous. Thick-walled exozonal growth and intrazoarial overgrowth with new zooecia budding from basal layer. Zooecial walls of subjacent growth show indistinct, broadly curved, orally convex lamination. Endozonal zooecia have granular cortex with thin, laminate zooecial linings. WT 178 Figure BULLETIN 291 EXPLANATION OF PLATE 9 1. Ceriopora micropora Goldfuss, 1826 .........22n-.2 eee All views are made from the lectotype, UB 119. This specimen was figured by Goldfuss, 1826, pl. 10, figs. 4a, d; von Hagenow, 1851, pl. 5, fig. 13 as Heteropora crassa; and Voight, 1953, pl. 2, fig. 4 as Pennipora beyrichi Hamm. als 1b. iG: 1d. Transverse section, acetate peel 200. Primary laminate structure arches convex orally; laminae appear to be con- tinuous across zooecial boundary zone. Intermediate dia- phragm has remnant laminar structure in planar oral portion, but remainder is completely replaced by clear cal- cite. Note relatively long, aborally flexed abutment which does not merge with zooecial wall. Transverse section, acetate peel 200; primary lamination partly preserved. Large, clear, calcite mass in oral monilis cuts across laminae sharply, and probably results from recrystallization of originally laminar tissue. Also, note intermediate diaphragm in zooecial cavity on right. Transverse section, acetate peel 200. Monili symmetrically to asymmetrically thickened and somewhat variably thick- ened longitudinally. Primary laminar structure is orally convex. Laminar structure grades aborally into indistinctly structured tissue whic hmay have originally been sublaminate to granular. Transverse section, acetate peel 100. Thick-walled exo- zonal growth shows irregularity in growth direction and variation in wall thickness. Note long tubelike (?) structure in central zooecial cavity. The structure merges with the zooecial wall at oral and aboral ends, and thus appears to be a primary structure. PLATE 9 BuLL. AMER. PALEONT., VOL. 69 PLATE 10 BULL. AMER. PALEONT., VOL. 69 am . CrERIOPpoRID CycLosToMEs (Bryozoa): NYE 179 EXPLANATION OF PLATE 10 Figure Page 1-3. Corymbopora menardi Michelin, 1846 2.0.0.0... 64 All specimens are from the Cenomanian, Le Mans (Sarthe), France. 1. MNHN Canu Coll. 57057-2 (1). la. Distal surface (capitula) of two branches X5 showing large polymorphs. 1b. Lateral view of branch shown above 5. Note distal ex- pansion forming capitula; small polymorphs are arranged in rows parallel to the growth axes of the branches. 1c. Detail of 1a X30 showing shape and arrangement of large dimorphs. 2. MNHN Canu Coll. 57057-2 (2). Surface of stem 30. Aper- tures of small polymorphs are arrayed in proximo-distal rows. The walls separating transversely adjacent zooecia form prominent ridges. 3. MNHN Canu Coll. 57057-2 (4). 3a, Transverse thin-section X30. Shows thin-walled large polymorphs in stem, and thick-walled small polymorphs covering outside of branch completely. 3b. Longitudinal thin-section X10 showing stem and expanded capitulum. Chamber in upper right portion of capitulum are lobes of two brood chambers. 3c. Tangential thin-section of stem X30. Section is slightly ob- lique. Small polymorphs are seen in the lower part of the figure, and a shallow, longitudinal section of large poly- morphs in the upper part of the figure. 3d. Longitudinal thin-section 30, detail of 3b. Cross sections of the lobes of two brood chambers can be seen. The posi- tions of two remnant, thick-walled stages are indicated by arrows. 4. MNHN Canu Coll. 57057-2 (3). 4a. Transverse thin-section of capitulum 30. Most zooecia exhibit thin-walled phase of growth. The section intersects a thick-walled phase in upper left portion of figure. 4b. Longitudinal thin-section of stem X30. Large polymorphs have thin, nearly parallel-sided walls with occasional inter- zooidal pores. Small polymorphs have short zooecial cham- bers and thick walls. 180 Figure 1-3. Corymbopora menardi Michelin, 1846 All BULLETIN 291 EXPLANATION OF PLATE 11 specimens are figured from the Cenomanien, Le Mans (Sarthe), France. il, la. 1b. lie: Nes we 3b. pw MNHN Canu Coll. 57057-2 (5). Longitudinal thin-section 10. Cavities in distal portion of capitulum are brood chambers. Longitudinal thin-section 30, detail of 1a. Note prominent interzooidal pores connecting longitudinally adjacent zooecial chamber of small dimorphs, and relatively large interzooidal pores connecting zooecial chamber of small and large dimorphs. Solid arrow is section through wall between trans- versely adjacent small polymorphs (ridge-forming wall) ; hollow arrow is section through wall between longitudinally adjacent small polymorphs. Longitudinal thin-section 30, detail of 1a. Shows large polymorph budding from distal wall of small dimorph. Longitudinal thin-section 100, detail of 1a showing struc- ture of wall between longitudinally adjacent zooecia. Laminae are indistinct but form broadly curved, orally con- vex patterns in this view. MNHN Canu Coll. 57057-2 (1) X400. Detail of Pl. 10, fig. 3c. Tangential section of small dimorphs. MNHN Canu Coll. 57057-2 (4). Transverse thin-section 100. Hollow arrow shows position of wall between transversely adjacent small polymorphs (ridge-forming) ; solid arrow shows position of wall be- tween longitudinally adjacent small polymorphs. Transverse thin-section, crossed nicols 400. Detail of 3a showing wall structure of small polymorphs. Laminae di- verge orally from the longitudinal boundary zone at a low angle in the portion of the wall between transversely ad- jacent zooecia. (The position of this portion of the wall is indicated by the hollow arrow in fig. 3a). The approximate position of the longitudinal boundary zone is indicated by two hollow triangles. Laminae are slightly arched, convex aborally in this view of the wall between longitudinally ad- jacent zooecia. (The position of this wall is indicated by the solid arrow in fig. 3a). Compare to the laminar configura- tion shown in fig. le which is at right angles to this view. BULL. AMER. PALEONT., VOL. 69 PLATE 11 PLATE 12 BULL. AMER. PALEONT., VOL. 69 + =») oor o . vi po a4 ce Figure CertoporIp CycLostomeEs (Bryozoa): Nye EXPLANATION OF PLATE 12 1,2. Corymbopora menardi Michelin, 1846 0 n.. All specimens are figured from the Cenomanien, Le Mans (Sarthe), France. ile MNHN Canu Coll. 57057-1. Distal surface of branch (capi- tulum) X30 showing brood chamber. Lobes of the brood chamber unite and are continuous with a single zooecial opening in lower portion of the figure. MNHN Canu Coll. 57057-2 (6). . Longitudinal thin-section X10. Shows two brood chambers and fasciculate appearance of autozooecia in stem and ex- panded capitulum. Note that expansion takes place as new, large polymorphs bud from the distal wall of small poly- morphs. Tangential thin-section 10 showing lobate profile of brood chamber. Detail of 2b X100. Detail of 2a 100 showing structure of more distal brood chamber. . Detail of 2a 100 showing lobe of more proximal brood chamber. 181 182 Figure BULLETIN 291 EXPLANATION OF PLATE 13 1. Coscinoecia radiata Canu and Lecointre, 1934 .......000000000 ee. The lectotype, MNHN Canu Coll. 58872-1, Miocene, Doué-la-Fon- taine (Marne-et-Loire), France, was figured by Canu and Lecointre, 1934, pl. 40, figs. 1-4. la. 1b. lic: 1d. le: ab lg. 1h. Zoarium 1. Surface of zoarium 5. Large polymorphs are arranged in rows radiating from the monticular areas. Small polymorphs are in monticules and in rows between large polymorphs. The monticule-like structure in the upper right portion of the figure is a brood chamber. Roof with overgrowth is par- tially broken away revealing hollow interior. Surface of zoarium X30. Monticule is shown in upper right. Large elliptical opening in lower left is continuous proxi- mally with a brood chamber. Longitudinal thin-section of growing tip X10 showing numerous, closely spaced, basal diaphragms in endozone. Longitudinal thin-section 10. Brood chamber in lower por- tion of figure. Primary growth is encrusted by a cheilo- stome bryozoan. The cheilostome is, in turn, encrusted by an intrazoarial overgrowth extending from a slightly more distal portion of the primary growth. Transverse section, acetate peel <5. Large cavities were made of boring organisms. The primary branch shows encrustation by a cheilostome bryozoan which is, in turn, encrusted from a more distal portion of the original zoarium. Longitudinal thin-section X30. Compare the relatively smooth-sided walls of large polymorphs (at about the center of the figure) to the variably thickened walls of the small polymorphs (at top of the figure). Basal diaphragms occur in zone of zooecial bending and in endozone. Tangential thin-section 30. Monticule is shown in right center. Compare pattern of zooecial arrangement to that seen in fig. 1b. The less distinct pattern in 1h is probably due to to the difficulty of discriminating between large small-poly- morphs and small large-polymorphs in tangential sections. BULL. AMER. PALEONT., VOL. 69 PLATE 13 a 24) 3 a t 4 MA D4 y pial pe) / ESE se Lg BULL. AMER. PALEONT., VOL. 69 PLATE 14 Figure Cerroporip CycLosTomeEs (Bryozoa): NYE EXPLANATION OF PLATE 14 1. Coscinoecia radiata Canu and Lecointre, 1934 ................ bet ae The lectotype, MNHN Canu Coll. 58872-1, Miocene, Doué-la-Fon- taine (Marne-et-Loire), France, was figured by Canu and Lecointre, 1934, pl. 40, figs. 1-4. la. 1b. lc. 1d. le. Ibe, lg. th. Longitudinal thin-section 100, detail of pl. 10, figs. le, g. Shows annularly thickened zooecial walls of small poly- morphs with alate monili; also, note broadly curved, orally convex laminae. Longitudinal thin-section 100. The zooecial wall of the large polymorph at lower right shows much less variation in thickness than the annularly thickened walls of small polymorphs. Tangential thin-section of intermonticular area 100. Note vague, pluglike bodies of light-colored tissue. Tangential thin-section of small polymorphs in monticular area X100. Note small, blunt, mural spines. Longitudinal thin-section of zooecial wall shared by small polymorphs 300. The cortex of most of the wall is com- posed of light-colored tissue with a dusting of small, dark grains. The structure is obscure, ranging from granular to faintly laminate. Distinctly laminate tissue caps, and partly surrounds, the cortex tissue. Longitudinal thin-section of large polymorph wall 300. Wall appearance is similar to le, but light-colored cortex tissue here forms discontinuous, pluglike bodies offset to one side of the wall. Tangential thin-section 300. Shows small polymorph with endozooecial spines to left and large polymorph to right. Note pluglike bodies of light-colored, indistinctly lami- nate cortex tissue bounded by distinctly laminated tissue. Tangential thin-section of large polymorph 300. Calcareous wall tissue is dusted with small, dark granules, and is broadly amalgamate. Note distinct lamination in outer wall, and small pluglike bodies in cortex. 183 184 Figure BULLETIN 291 EXPLANATION OF PLATE 15 1. Coscinoecia radiata Canu and Lecointre, 1934 ......00.0.00. The lectotype, MNHN Canu Coll. 58872, Miocene, Doué-la-Fon- taine (Marne-et-Loire), France, was figured by Canu and Lecointre, 1934, pl. 40, figs. 1-4. tas 1b. lc. 1d. le. 16 1g. 1h. Longitudinal thin-section, detail of pl. 10, fig. le, 100. Zooecia in intrazoarial overgrowth directed proximally in re- lation to major grewth axis of branch. Note intermediate dia- phragms and thin basal layer. Longitudinal thin-section 100. Shows basal diaphragms in outer endozone, zooecial growth directed towards upper left. Transverse thin-section 30. Large cavity is brood chamber; note incomplete floor. Longitudinal thin-section, detail of 1f 300. Shows brood chamber roof and zooecial wall of small polymorph grow- ing to the left. Tangential thin-section 30. Large cavity is a brood chamber. Longitudinal thin-section 3. Large cavity is a brood chamber. Longitudinal thin-section of zooecial wall 300. Calcareous microstructure is laminate except for a small mass of granu- lar tissue in aboral part of cortex. Longitudinal thin-section, detail of 1f 300. Shows cavity of brood chamber to left, cavity of subjacent zooecia to the right. Lateral growth from oral tips of the subjacent zoo- ecial wall was incomplete. The gap was later sealed by the emplacement of a thick, indistinctly laminate, intermediate diaphragm. . Longitudinal thin-section of a basal diaphragm 300. BuLL. AMER. PALEONT., VOL. 69 PLATE 15 BULL. AMER. PALEONT., VOL. 69 PLATE 16 Figure CERIOPoRID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 16 1. Diplocava incondita Canu and Bassler, 1926 .......0.0.0..0000..000000... The lectotype, USNM 69925-2, Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland, was figured by Canu and Bassler, 1926, pl. 10, fig. 5 (lower right), and 6. ae 1b. 1c. 1d. le. Af: lg. ih. Zoarial fragment <5. Shows irregular growth habit. Surface of zoarial fragment X30. Shows intrazoarial over- growth on upper right. Tangential thin-section X30. Note narrowly integrate ap- pearance of zooecial walls. Longitudinal thin-section X30. Much of the primary wall structure altered by diagenetic processes. Tangential thin-section 30. Section somewhat deeper than those figured in 1c, f, h. Zooecial walls are amalgamate in this section. Tangential thin-section 100. Zooecial walls show both in- tegrate and amalgamate appearance. Longitudinal thin-section 100. Laminae directed orally from boundary zone, broadly recurve aborally and adjoin thin zooecial lining. Apertural rim of zooecial wall is structurally continuous with simple external wall. Also note thin basal diaphragm. Thin-section transverse to zoarium 30. Zooecia in center have integrate appearance. Note that most zooecia in over- growth grow relatively straight from basal layer and are thick-walled. 185 186 Figure 1-6. Diplocava incondita Canu and Bassler, 1926 BULLETIN 291 EXPLANATION OF PLATE 17 All specimens from Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland. te 5b. 5c. 6a. 6b. USNM Loc. 2404-3, longitudinal thin-section 30. Shows growth habit with long, continuously growing zooecia uncom- plicated by intrazoarial overgrowth. USNM Loc. 2404-1, tangential thin-section X30. Section passes through a simple external wall in upper right portion of figure. USNM Loc. 2404-8, longitudinal thin-section 30. Shows mode of growth typified by repetitive overgrowth. Zooecia are initially thick-walled, then slightly thinner, and straight. Two basal diaphragms are seen in third zooecium from bottom. USNM Loc. 2404-5, tangential thin-section 30. Shallow tangential section in upper right portion of figure shows zooecial walls with integrate appearance. Deep tangential section of intrazoarial overgrowth in lower left portion of figure. USNM Loc. 2404-9. Longitudinal thin-section 30. Longitudinal thin-section 100. Shows thick external wall and intrazoarial overgrowth. Longitudinal thin-section 100. Zooecial chamber continuous from subjacent growth to overgrowth. Zooecial wall on right apparently grew continuously from subjacent growth to overgrowth, but growth on left was discontinuous. Lectotype 69925-2, figured by Canu and Bassler, 1926, pl. 10. fig. 5 (lower right). Transverse thin-section 100. Shows extension of zooecial lining over aboral surface of simple external wall. Longitudinal thin-section 100. Zooecial walls show faint recurved lamination. Note thick zooecial linings in zooecia recumbent upon basal layer. Dark line separates two growth episodes. PLATE 17 BuLuL. AMER. PALEONT., VOL. 69 re eS) re PLATE 18 BULL. AMER. PALEONT., VOL. 69 s a Tai ‘ _ dq, AF 7 << Bae > ay G Fem tb “oF i rs ta * z Figure Crer1oporIp CycLosToMEs (Bryozoa): Nye EXPLANATION OF PLATE 18 1-3. Diplocava incondita Canu and Bassler, 1926 ............00000e. All specimens figured from Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland. 1 USNM Loc. 2404-6. Transverse section, acetate peel 200. Zooecia on left grew orally towards right, have simple external walls, and are encrusted on right by intrazoarial overgrowth. Basal layer and most of wall of overgrowth silicified. Pores through simple external wall in lower zoo- ecium sealed aborally by laminate calcareous tissue con- tinuous with zooecial lining. Note narrow extension of zooecial wall to basal layer of overgrowth. Apertural opening of zooecium in upper left part of figure sealed by basal layer of overgrowth. Lectotype, USNM 69925-2. Longitudinal section, acetate peel X200. Zooecial wall shows laminae arching orally from boundary zone and recurving aborally. Note apertural tip of zooecial wall extending to basal layer (replaced by silica) of intrazoarial overgrowth. Simple external wall to left separated from zooecial wall by long pore extending obiquely to surface of diaphragm. Pseudopores of simple ex- ternal wall are sealed aborally by laminate calcareous tissue continuous aborally with zooecial lining and extending orally to form peristome. Also, note thin (one lamina) basal layers in more aboral portions of zooecial chambers. USNM Loc. 2404-7. Tangential section, acetate peel 200. Zooecial walls have narrowly integrate appearance. Cal- careous tissue of peristomial diaphragms merges continuously with zooecial wall. Note laminar structure of peristome (lineations) concentric with restricted apertures in simple ex- ternal wall. 187 188 Figure BuLLETIN 291 EXPLANATION OF PLATE 19 1-3. Diplocava incondita Canu and Bassler, 1926 .........0000000000.0.... All specimens are figured from Cretaceous, Valangian, Ste. Croix, Switzerland. 1. USNM Loc. 2404-7. Tangential section, acetate peel 200. Zooecial walls have variously developed integrate ap- pearance. Note deflection of thick zooecial lining at inter- zooidal pore connecting central and distal zooecial chambers. USNM Loc. 2404-5. Longitudinal section, acetate peel 200. Zooecia on right grew orally towards left side of figure and have thick, laminate, intermediate diaphragms near aperture. Diaphragms have thick, aborally flexed abutments. USNM Loc. 2404-1. Longitudinal section, acetate peel 500. Apertural portion of zooecial wall and oblique section of simple external wall (with circular pores) in lower part of figure overlain by intrazoarial overgrowth. Basal layer and most of proximal zooecial wall of intrazoarial overgrowth is replaced by silica. In subjacent zoarial growth unit, calcareous tissue of zooecia] wall is structurally continuous with calcareous tissue of simple external wall (marked by pores). Also, note narrow extension of zooecial wall to basal layer of overgrowth in center of figure marking approximate position of zooecial boundary zone. BULL. AMER. PALEONT., VOL. 69 PLATE 19 \ Wee ‘ vo. ve 5 aw vires a ye os, Be et ger é “oe e “L = Ba wa PLATE 20 BULL. AMER. PALEONT., VOL. 69 ae CERIoporID CycLosToMEs (Bryozoa): NYE 189 EXPLANATION OF PLATE 20 Figure Page 1. Ditaxia anomalopora (Goldfuss), 1826 20000000000 eee. 88 Lectotype, UB 120, was figured by Goldfuss, 1826, pl. 10, figs. 5c, d; and by von Hagenow, 1851, pl. 4, fig. 9c. Cretaceous, Maastrichtian, Petersberg near Maastricht (Limburg), Nether- lands. la. Zoarial surface X30. Shows monticular areas and irregular distribution of large and small polymorphs in intermonticu- lar areas. 1b. Zoarial fragment <5. The zoarium is subcylindrical in proximal portion, expanding to frondose distally. Note patchlike distribution of monticules. lc. Tangential thin-section 30. 1d. Longitudinal thin-section 30. Shows bifoliate growth habit; median lamina has dark line at medial boundary zone. le. Transverse thin-section 30. 190 Figure BULLETIN 291 EXPLANATION OF PLATE 21 1,2. Ditaxia anomalopora (Goldfuss), 1826 22.000....0...0cccccecccceeeeees Cretaceous, Maastrichtian, Guelem, Maastricht (Limburg), Nether- lands. ls 2b. 2c. USNM Loc. 2404-5. Longitudinal section, acetate peel 100. Shows median layer and recumbent zooecia. Lineations diverging distally from median plane of median layer are interpreted as remnants of primary lamination. USNM Loc. 2404-2. Longitudinal section, acetate peel 100. Shows intermediate diaphragms in large polymorph and in small polymorph just distal to it. Longitudinal section, acetate peel 100. Large polymorph at top of figure is nearly parallel-sided, but lunarial tissue in proximal wall is obscure because of recrystallization. Proxi- mally, small polymorphs show prominent annular thickenings. Slightly oblique longitudinal section, acetate peel «50. Shows intermediate diaphragms in large polymorph at top and center right part of figure. Thick-walled growth in exozone is followed by thin-walled zooecia] growth, and a change in zooecial growth orientation suggests a rejuvenated growth phase. Interzooidal spaces, apparently sealed orally and aborally by thin, calcareous walls, are seen between large polymorphs in rejuvenated zone. BuLL. AMER. PALEONT., VOL. 69 PLATE 21 BULL. AMER. PALEONT., VOL. 69 PLATE 22 Figure CERIOPORID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 22 1,2. Ditaxia anomalopora (Goldfuss), 1826 ......0....00000000000 000 ee Cretaceous, Maastrichtian, Guelem, Maastricht (Limburg), Nether- lands. All photographs taken from acetate peels. i 2b. Ze: USNM Loc. 2405-6. Longitudinal section 100. Walls of small polymorphs show large increase in thickness from endozone to exozone. In exozone, walls of all small poly- morphs in figure show annular thickening at approximately same level. USNM Loc. 2405-5. Longitudinal section 100. Large polymorph has thin, near- ly parallel-sided walls, although distal walls show some variation in thickness most noticeable just oral to zooecial flexure. Proximal wall of large polymorph has homogeneous calcareous tissue making up a lunaria-like structure. Longitudinal section 100. Large polymorph with thin, nearly smooth-sided zooecial wall and lunaria-like structure in proximal wall. Zooecial walls of small polymorphs are thickened annularly. Detail of 2a showing wall structure of large polymorph. Proximal wall has homogeneous tissue forming lunaria-like structure. Laminae in outer cortex of adjacent small poly- morph dip steeply in aboral direction. 191 192 Figure BULLETIN 291 EXPLANATION OF PLATE 23 1. Haploecia straminea (Phillips), 1829 .00....0000occccceceecceeceesenteeeee Lectotype, YM-T81/2, Jurassic, Bajocien (?), Bathonien (?), Yorkshire, England. The thin-sections figured in lc-f were made from unencrusted branches at less than 5 mm from the growing tip. la. 1b. lc: 1d. le. like Nearly complete zoarium 2 showing ovoid outline of branching colony. Surface of branch X50. Shows newly-formed simple ex- ternal walls close to the growing tip. The surfaces of the external walls are slightly depressed relative to the thin apertural parts of the zooecial wall, and zooecia have poly- gonal, generally hexagonal, outlines. Longitudinal thin-section 100. Shows newly-formed simple external wall. Wall is thin and delicate in appearance; com- pare to more robust appearance of external wall in pl. 24, fig. 1f or pl. 26, figs. la, c, d. Zooecial wall shows slight asymmetry in thickness across zcoecial boundary zone and thin zooecial linings. Zooecial wall in right center shows remnant laminae arched orally convex. Longitudinal thin-section 50. Shows zone just distal to growing tip where simple external walls first appear. Tangential thin-section 100. Shows amalgamate appear- ance of laminate zooecial walls; also shows interzooidal pores. Transverse thin-section X50. Shows central zooecium dis- tinctly larger in diameter than surrounding zooecial open- ings. PLATE 23 BULL. AMER. PALEONT., VOL. 69 PLATE 24 BuLL. AMER. PALEONT., VOL. 69 % s ae 4 tz aT) ® coo rey t ys Figure CeRIoporID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 24 1. Haploecia straminea (Phillips), 1829 2000000000oooccccocecceeccceeeeees Paralectotype YM 1T81/1, Jurassic, Bajocien (?), Bathonien (?), Yorkshire, England. la. 1b. lc. 1d. le. 1f. Longitudinal thin-section 30 showing abandoned growing tip and intrazoarial overgrowth. Zooecia in abandoned branch tip are thick-walled and have simple external walls. Transverse thin-section X30 showing thick walls in exozone, thin walls in endozone. At top center, peristome protrudes into overgrowth. Longitudinal thin-section X30 showing narrow zooecia in endozone with diameters widening distally in exozone. Zooecia of primary branch have simple external walls. Tangential thin-section 100. Large zooecium at center- left shows interzooidal pore to left and more closely-spaced pseudopores in terminal diaphragm to right. Tangential thin-section 100 showing simple external walls. Longitudinal thin-section 100 showing structure of zoo- ecial wall in exozone and simple external wall. Boundary with thin basal layer of overgrowth is marked by a distinct, dark line. 193 194 Figure BULLETIN 291 EXPLANATION OF PLATE 25 1,2. Haploecia straminea (Phillips), 1829 ...................ccccccccccceesseeeesteeree Both specimens from Jurassic, Bajocien (?), Bathonien (?), Yorkshire, England. All photographs taken from acetate peels. 1. la. 1b. Ic. 1d. 2b. YM T81/1, paralectotype. Intersection of two different branches 50. Curved line from top left to lower right side of figure marks intersec- tion between distally growing branch below (longitudinal section) and side of branch above (transverse section). Complex growth pattern in lower right is an intrazoarial overgrowth of branch to left. Zooecia in branch to upper right had thick simple external walls apparently in place be- fore intersection occurred. Thin, zooecial seals of left-hand branch were apparently deposited at about the same time as intersection occurred (detail in figs. 1c and 1d). Longitudinal section 200. Shows basal diaphragms in inner exozone. Only distal-most diaphragm is well preserved. Diaphragms are flexed slightly in oral direction, but ap- pear to abut, rather than merge with, zooecial wall. Detail of 1a 200. Shows boundary between intersecting branches (indicated by arrow). Zooecium on right is sealed by a simple external wall. Thin-walled zooecia on left are sealed by nonporous diaphragms composed of light-colored, granular tissue (directly adjacent to plane of intersection) continuous with cortex of zooecial wall, and lined aborally by laminated tissue continuous with zooecial lining. Detail of 1a 200 showing boundary between intersecting branches (indicated by arrow). Thick-walled zooecium on left is sealed by a thin diaphragm which flexes aborally to merge with zooecial! lining. Faint line and thin gap separate both branches. Zooecium on right has a thick, poorly-pre- served simple external wall. YM 1T81/2, lectotype. Longitudinal section near growing tip 200. Thin, newly- formed external wall merges continuously with calcareous tissue of zooecial wall. Longitudinal section near growing tip 200. Shows thin, newly-formed simple external wall. Calcareous tissue of wall flexes aborally to merge continuously with zooecial wall. BULL. AMER. PALEONT., VOL. 69 PLATE 25 BULL. AMER. PALEONT., VOL. 69 PLATE 26 Figure 1. Haploecia straminea (Phillips), 1829 000000000000 Crerioporip CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 26 Paralectotype from Jurassic, Bajocien (?), Bathonien (?), York- shire, England. All photographs taken from acetate peels. la. 1b. lc. 1d. Longitudinal section 200. Shows single zooecium with simple external wall on left, separated by a dark line (in- dicated by an arrow) from intrazoarial overgrowth in transverse section on far right. Calcareous tissue of simple external wall is light-colored (probably granular) just sub- jacent to dark boundary line, but distinctly laminate in aboral part; laminae are moderately convex in aboral direc- tion. Basal layer and cortex of overgrowth have light- colored, homogeneous calcareous tissue. Note thick, distinctly laminate zooecial lining. Transverse section of branch on bottom 100. Shows oblique longitudinal sections of intrazoarial overgrowths, each marked by simple external walls. Note basal diaphragm in peristome of lower left-hand zooecium, and intermediate diaphragm abutting simple external wall in zooecium at middle of figure. Poorly preserved lineations of cortex tis- sue in zooecial walls of bottom branch are interpreted as original laminations directed convex orally. Longitudinal section 200. Zooecium on right with simple external wall (peristome directed obliquely towards upper left corner) is separated by two dark lines and a gap from intrazoarial encrustation to left (indicated by arrow). Cal- careous tissue of peristome merges continuously with sub- jacent zooecial wall. The peristome and external wall have a laminate inner portion continuous with zooecial lining, and light-colored, subgranular outer portion continuous with the cortex of the zooecial wall. Longitudinal section on left and obliquely transverse section of intrazoarial overgrowth on right 200. Peristomes of zooecia on left are directed almost horizontally to right. External walls are continuous structurally, with calcareous tissue of subjacent zooecial walls. Dark jine marks boundary between growth phases. 195 Page 98 196 Figure BULLETIN 291 EXPLANATION OF PLATE 27 1-3. Haploecia multilamellosa (Canu and Bassler), 1926 ...................... All specimens figured were identified by R. S. Bassler and were collected from the type locality in the Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland. ik la. 1b. Ic. 1d. Se: 3d. Lectotype, USNM 69922-1, figured by Canu and Bassler, 1926, pl. 9, figs. 1 (center specimen), 5, 6. Most of the in- ternal structure is obscured by recrystallization and silicifica- tion. Branch X5. Surface of branch X30. Zooecia are arranged in rows; zooecial apertures in adjacent rows alternate in position. Peristomes are commonly located centrally with respect to the zooecial walls. Longitudinal section, acetate peel X30. Zooecia intersect zoarial surface obliquely; zooecia encrusting primary branch grow orally in approximately the same direction as under- lying zooecia. Tangential section, acetate peel X30. Zooecial openings are commonly elliptical, with greatest dimension parallel to the growth axis of the branch. Transverse section, acetate peel X30. Shows primary branch and single intrazoarial overgrowth. A _ single, enlarged zooecium occupies the center of the endozone. Paralectotype, USNM 69922-2. Branch 5. Surface of branch X30. Arrangement of apertures are not as regular as those seen in PI. 27, fig. la. Topotype identified by R. S. Bassler, USNM 2384-1. Longitudinal section, acetate peel 30. Shows growing tip of primary branch with a thin, intrazoarial overgrowth. Tangential section, acetate peel X30. Tangential to zooecia in primary branch, shows shape and arrangement of zoo- ecial openings. Transverse section, acetate peel 30. Shows primary branch with enlarged central zooecium and three generations of intrazoarial overgrowth (upper right). All zooecia, except in outer-most overgrowth, have simple external walls. Tangential section, acetate peel X30. Section is tangential to primary branch in axial portion, and longitudinally oblique to intrazoarial overgrowth on either side. Zooecial apertures are arranged in longitudinal rows; zooecia in adjacent rows generally alternate in position longitudinally. Smaller round opening at right center is a peristome. PLATE 27 BuLL. AMER. PALEONT., VOL. 69 PLATE 28 BULL. AMER. PALEONT., VOL. 69 ee Figure CERIoporID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 28 1-3. Haploecia multilamellosa (Canu and Bassler), 1926 .........000.0........ All specimens figured were identified by R. S. Bassler, and were collected from the type locality in the Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland. ile iia 1b. le; 1d. le. 3b. USNM Loc. 2384-2. Longitudinal section, acetate peel 30. Shows primary branch with coaxial endozone and exozone enveloped by two intrazoarial overgrowths. Zooecium in upper portion of figure has a prominent peristome submerged beneath basal layer of overgrowth. Tangential section, acetate peel X30. Section intersects a primary branch (upper right) and an intrazoarial over- growth. In the lower portion of the figure, two circular peri- stomes from the subjacent primary branch extend into the overgrowth. Tangential section, acetate peel 100. Shows pseudopores and restricted aperture in simple external wall. Section is slightly oblique; !ower portion of figure shows thick, exo- zonal zocecial walls of primary branch; upper part of figure shows thin endozonal walls of intrazoarial over- growth. Longitudinal section, acetate peel 100. Detail of 1a show- ing wall structure. Note thin band of light-colored tissue in outer cortex which marks the zooecial boundary zone, and the rodlike extensions from the cortex forming prominent, intrazooecial spines. Zooecia of the primary growth are sealed by simple external walls, and are encrusted by the basal layer of intrazoarial overgrowth. A thin, light-colored boundary zone separates the two (detail, Pl. 31, fig. 2b). Tangential section, acetate pee] 100. Section is just aboral to the simple external wall. Note the prominent mural spines extending from the cortex. The spines are partially covered by zooecial lining. Interzooidal pore shown in lower left is narrow, with little flare at either end. USNM Loc. 2384-1. Transverse section, acetate pee] 100. Shows the large central zooecium which lacks zooecial lining. Smaller surrounding zooecia typically have a thin zooecial lining. In exozone, zooecial walls have a few small, mural spines; zooecia are sealed by simple external walls which merge continuously with the zooecial lining. USNM Loc. 2384-10. Tangential section, acetate peel 100. Shows typical ar- rangement of zooecia and shape of openings. Note thick zooecial lining which covers most intrazooecial spines com- pletely. The spines appear as light-colored rods contrasting with the dark-colored, longitudinaly laminated tissue of the zooecial lining. Longitudinal section, acetate peel 100. Shows primary branch and intrazoarial overgrowth along left side of figure. In primary branch, zooecia grow towards upper left and have simple external walls. In the lower left zooecium, the peristome is low, and the peristomial aperture is sealed by calcareous tissue continuous with the simple external wall. The basal layer of the overgrowth is poorly differentiated from subjacent calcareous tissue of the primary branch; but, generally, a thin, light-colored boundary zone separates the two. 197 198 Figure BULLETIN 291 EXPLANATION OF PLATE 29 1-3. Haploecia muitilamellosa (Canu and Bassler), 1926 ...................... All specimens were identified by R. S. Bassler and were collected from the type locality in the Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland. ile ila 1b. Ne. 1d. le. 2b. Zc; USNM Loc. 2384-10. Longitudinal section, acetate peel 5. Shows intersection of a distally growing branch with a second branch. The plane of intersection is visible as a dark line (figs. 1c, 1d for detail). Note that the intersection occurred before either primary branch was covered by an intrazoarial overgowth. The overgrowth on the left-hand branch partially envelops the intersecting branch. Tangential section, acetate peel X30; at intersection shown in la. Plane of intersection appears as a dark line. Over- growth zooecia are visible in upper right. Longitudinal section, acetate peel 30, detail of 1a. Shows primary branch in center, intrazoarial overgrowth to the left, and intersecting primary branch on right. Zooecial de- tails of intersecting branch are obscure because of silicifica- tion of the zooecial walls. Longitudinal section, acetate peel 30; detail of Jc. Dia- phragms were deposited by zooids on both sides of the zone of intersection. To the left are simple external walls. Dia- phragms on the right are thin and non-porous. Longitudinal section, acetate peel 100 showing thin, inter- mediate diaphragms in intrazoarial overgrowth. USNM Loc. 2384-8. Longitudinal section, acetate peel 5 showing oblique inter- section of two distally growing branches. In contrast to 1a, the intersection apparently occurred at the actively growing tips of both branches. Note the deflection of endozonal zooecia so that the major axis of distal growth of each branch assumes the approximate orientation of the other branch. Details are shown in 2b and 2c. Detail of 2a X30. Proximal zooecia are sealed off at the zone of intersection (see 2c). Distally, zooecia merge con- tinuously along the zone of intersection and curve away from the zone of intersection forming a common endozone for a short distance. Detail of 2b 100 showing closure of zooecia along the zone of intersection. Note the slight distal flexure and thinning of zooecial walls just aboral to the zone of intersection. USNM Loc. 2384-1. Longitudinal section, acetate peel 100. Shows primary growth on lower left and intrazoarial over- growth on the upper right. Zooecial wall of the overgrowth have light-colored (interpreted as granular) cortices and thin, dark-colored zooecial linings which merge continu- ously with the basal layer. The basal layer is difficult to separate from the encrusted zooecial wall and peristomial diaphragm of the primary branch, but a thin, light-colored zone generally marks the boundary. PLATE 29 BULL. AMER. PALEONT., VOL. 69 PLATE 30 AMER. PALEONT., VOL. 69 BULL. Figure CERIOPpoRID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 30 1-3. Haploecia multilamellosa (Canu and Bassler), 1926 ...........0.00.0..... All specimens figured were identified by R. S. Bassler and were collected from the type locality in the Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland. ils la. 1b. USNM Loc. 2384-2. Transverse section, acetate peel 200. Shows primary branch and two consecutive intrazoarial overgrowths. Over- growths are one and one-half to two zooecial diameters thick. Intrazoarial overgrowths are separated from each other and from the primary branch by a dark line or by a thin, light-colored boundary zone (indicated by arrows). The simple external wal] of the zooecium in the center of the figure is thin with dark Jaminae similar in thickness and appearance to zooecial lining. The wall is apparently con- tinuous with the zooecial lining to left, but separated on right by a pseudopore. Wall is laminate; basal layer of suprajacent overgrowth is light-colored with indistinct struc- ture interpreted as granular. Longitudinal section, acetate peel 200. Shows primary branch on right and two intrazoaria] overgrowths, each separated by a thin, light-colored boundary zone (indicated by arrows). Zooecia grew towards upper left. Zooecial aper- tures are sealed by simple external walls. USNM Loc. 2384-7. Longitudinal section, acetate peel 50. Shows interruption of zooecial growth in endozone just below center of figure, and renewed growth indicated by offset of zooecial walls. Also note variation in size of central zooecium in lower portion of figure; diameters of other endozonal zooecia show relatively constant diameter. USNM Loc. 2384-10 200. A thin, intermediate diaphragm is shown in the central zooecium of the intrazoarial over- growth. The diaphragm shows a slight aboral flexure at juncture with zooecial wall. Primary branch is seen along right side of figure with intrazoarial overgrowth to the left. The primary branch and the basal layer are separated by a thin, light-colored zone (indicated by arrow). Note the small intrazooecia! spines in the basal layer. Note slightly undulatory growth and occasional small mural spines typical of endozonal appearance; also, the relatively large inter- zooidal pore in the zooecial wall at lower center. 199 200 Figure BULLETIN 291 EXPLANATION OF PLATE 31 1,2. Haploecia multilamellosa (Canu and Bassler), 1926 .................... All specimens were identified by R. S. Bassler and collected from the type locality in the Cretaceous, Valangian, Ste. Croix (Vaud), Switzerland. il la. 1b. Zia 2b. USNM Loc. 2384-10. Transverse section, acetate peel 200. Shows proximal growth separated from overgrowth by a dark line and, sometimes, by a thin, light-colored boundary zone (indicated by arrow). Simple external wall of zooecium in proximal growth is thin and composed of dark laminate tissue which recurves aborally to merge with zooecial lining, contrasting with lighter-colored and homogeneous calcareous tissue of superposed zooecial walls interpreted here as originally granular. Note thin, dark-colored zooecial linings of recum- bent overgrowth zooecia. Longitudinal] section, acetate peel 200. Shows oral growth of zooecium on right directed toward left-hand side of figure. Zooecium is capped by a simple external wall and encrusted by intrazoarial overgrowth. Zooecia in overgrowth grow orally towards upper left. Laminate tissue of simple external wall flexes aborally to merge with zooecial lining; note oblique section of pseudopore in proximal portion of external wall which appears to separate external wall from compound zooecial wall. Recumbent zooecial walls of over- growth are very thin, and are separated from proximal growth by a thin, light-colored zone (indicated by arrow). USNM Loc. 2384-2. Longitudinal section, acetate peel 500. Shows apertural portion of zooecium on right with simple external wall (middle of figure), and recumbent zooecia of intrazoarial overgrowth on left. Aperture in peristome is sealed by laminate calcareous tissue merging continuously with cal- careous tissue of external wall. Recumbent wall of the over- growth zooecium is separated from simple external wall by thin, light-colored zone (indicated by arrow). Longitudinal section, acetate peel 500. Detail of Pl. 28, fig. 1d. Zooecia on left grew orally toward right side of figure, capped by simple external wall (from center top to center bottom of figure) and encrusted by intrazoarial over- growth. The zooecium in the overgrowth grew orally towards top of figure. The overgrowth is separated from the subjacent zooecial wall and peristomial diaphragm by a thin, light-colored boundary (indicated by arrow). Note the prominent mural spines in the zooecial wall to the left. The spines have cores composed of homogeneous-appearing calcareous tissue, and are generally submerged beneath laminated zooecial lining. A few light-colored cores of small mural spines can be seen in the wall of the recumbent zoo- ecium on the right; the cores are nearly completely sub- merged beneath the zooecial lining. PLATE 31 BULL. AMER. PALEONT., VOL. 69 a tl tala esp x f. ee Dap. Oe ete PLATE 32 BuLL. AMER. PALEONT., VOL. 69 Figure CeRIoporID CycLosTomEs (Bryozoa): NYE EXPLANATION OF PLATE 32 1. Heteropora cryptopora (Goldfuss), 1826 .0.........0..0ccccccccccccceecceet ees Lectotype, UB 118a, figured by Goldfuss, 1826, pl. 10, fig. 3a; by von Hagenow, 1851, pl. 5, fig 6; and by Canu and Bassler, 1920, text fig. 222A, p. 681. From Cretaceous, Maastrichtian, St. Petersberg near Maastricht (Limburg), Netherlands. la. 1b. Ic. 1d. le; lite Zoarium X2. Zoarium is massive with bulbous to digitate branches. Surface of zoarium X30. Shows zooecial apertures with nearly continuous variation in size. Tangential thin-section 30. Shows zooecial openings which are commonly elliptical to subelliptical, and have nearly con- tinuous variation in size. Transverse thin-section 30. A dark line separates zoarial growth phases. Some zooecial chambers subjacent to over- growth are filled with dark-colored, fine-grained sediment; other zooecial chambers are filled with clear calcite. Longitudinal thin-section X10. Shows zoarial growth by repetitive addition of superposed intrazoarial overgrowths. Chambers in lower right and upper left are brood cham- bers; both are located proximally with respect to individual growth phases. Longitudinal thin-section 30, detail of le. Shows struc- ture of brood chamber. In this view, five zooecia pass through chamber to roof. Intermediate diaphragms are seen in many zooecia suboral to basal layer of overgrowth. Two intermediate diaphragms can also be seen in lower right portion of figure. 201 202 Figure BULLETIN 291 EXPLANATION OF PLATE 33 1,2. WHeteropora cryptopora (Goldfuss), 1826 00.0.0... Both specimens figured were collected from the Cretaceous, Maastrichtian, St. Petersberg near Maastricht (Limburg), Netherlands. 1. Lectotype, UB 118a, figured by Goldfuss, 1826, pl. 10, fig. Za: 2b. 2c. 3a; by von Hagenow, 1851, pl. 5, fig. 6; and by Canu and Bassler, 1920, text fig. 222A, p. 681. Longitudinal thin-section 100 showing structure of brood chamber. Zooecia and septate structures pass through the chamber. These intra- chamber structures have thin, dark-colored walls in contrast to the lighter-colored tissue of the walls distal to the brood chamber. Septate structure on upper left is continuous with intra-chamber zooecium in oblique section. The roof is porous and separated from the superjacent encrusting growth by a thin, dark line. Paralectotype, UB 118b, figured by Goldfuss, pl. 10, fig. 3b. Longitudinal thin-section 100 showing characteristic ap- pearance of zooecial wall. Thin intermediate diaphragms are shown just aboral to apertures of zooecia subjacent to overgrowth phase on right. Longitudinal thin-section 10 showing characteristic ap- pearance of zoarial growth by repetitious increments of in- trazoarial overgrowths. The outline at the termina] sur- face of each unit is emphasized by distally convex, dark lines caused by emplacement of intermediate diaphragms roughly coincidental with neighboring zooecia, and by em- placement of a thin, dark basal layer over the subjacent zoarial surface. Branching begins with the development of two separate, but later confluent, growth units on the same zoarial surface. Longitudinal thin-section 30, detail of 2b. PLATE 33 BULL. AMER. PALEONT., VOL. 69 PLATE 34 BULL. AMER. PALEONT., VOL. 69 Figure CERIOPORID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 34 1,2. Heteropora cryptopora (Goldfuss), 1826 2.0.0.0... 1. la. 1b. 2a. 2b. Paralectotype, UB 118b, from the Cretaceous, Maastrich- tian, St. Petersberg near Maastricht (Limburg), Nether- lands; figured by Goldfuss, 1826, pl. 10, fig. 3b. Longitudinal section, acetate peel 100 showing charac- teristic profile of apertural parts of zooecial walls. Inter- mediate diaphragms occur at nearly the same level in all zooecia, slightly aboral to the terminal growth surface. The basal layer of the encrusting growth unit is thin and dark in color. This view of the overgrowth shows the thin-walled, parallel-sided appearance typical of zooecia in the endozone. Transverse thin-section X30. Growth phases are separated by two dark lines. The line on the right is caused by inter- mediate diaphragms; the line on the left is thin, dark- colored basal layer. Specimen USNM Loc. 2387-11 from the Cretaceous, Maastrichtian, near Maastricht (Limburg), Netherlands. Tangential thin-section 30. Shows section cutting two in- trazoarial growth phases. Surface of juncture is about the middle of figure. Longitudinal thin-section 30. Shows parts of three over- growth phases. Large cavity in proximal portion of middle unit is a broad chamber. In middle zoarial growth phase, intermediate diaphragms occur just aboral to terminal growth surface. Also, a few intermediate diaphragms are scattered throughout the growth unit. 203 204 Figure BuLLeTIN 291 EXPLANATION OF PLATE 35 1,2. Heteropora cryptopora (Goldfuss), 1826 000000000... Both specimens figured were collected from the Cretaceous, Maastrichtian, St. Petersberg near Maastricht (Limburg), Netherlands. 1. la. 1b. lc. 2a. 2b. Paralectotype, UB 118b, figured by Goldfuss, 1826, pl. 10, fig. 3b. Longitudinal section, acetate peel 200. Diaphragm in zoo- ecial chamber on right shows slight aboral flexure and is interpreted as an intermediate diaphragm. Diaphragm in left-hand zooecium shows slight oral flexure at juncture with zooecial wall, but appearance may be due to recrystalliza- tion rather than primary structure. Zooecial chamber and zooecial wall to center right are continuous with suprajacent growth phase. Longitudinal section, acetate peel 200. Shows distal por- tion of one growth unit and proximal portion of supra- jacent intrazoarial overgrowth. Zooecial walls thin sym- metrically near aperture. Note distinctly laminate basal layer of overgrowth, draping over apertural parts of zoo- ecial wall. Poorly preserved intermediate diaphragm is shown in central zooecial chamber; diaphragm has short, aborally flexed abutments. Longitudinal section, acetate peel 200. Basal layer is un- dulatory, extends diagonally from upper right to lower left side of figure, and separates subjacent from suprajacent in- trazoarial overgrowth (dark arrow). Intermediate dia- phragms occur in zooecial chambers of subjacent growth unit (hollow arrows). Diaphragm in right center zooecium is dark in color and indistinctly laminate. Undulatory basal layer forms recumbent wall of most proximal overgrowth zooecium. More distal wall has optically structureless (orig- inally granular?) calcareous tissue and thin zooecial lining. Lectotype, UB 118a, was figured by Goldfuss, 1826, pl. 10, fig. 3a; by von Hagenow, 1851, pl. 5, fig. 6; and by Canu and Bassler, 1920, text fig. 222A, p. 681. Longitudinal section, acetate peel 200. Shows primary laminate structure of exozonal zooecial wall. Laminae are convex orally and continuous across zooecial boundary zone. Patches of homogeneous calcareous tissue occur in more aboral parts of outer cortex. Longitudinal section, acetate peel 500. Zooecia grew orally to left. Shows apertural portion of zooecial wall, with zone of irregular structure at about center of picture making boundary between subjacent growth phase and suprajacent overgrowth phase. Zone of irregular structure is symmetri- cal with basal layer to either side. Zooecial chamber is con- tinuous into overgrowth zooecium, but zooecial wall shows marked thinning. Also, note short stublike projection of zooecial wall in center part of figure (indicated by arrow), perhaps a remnant of a resorbed zooecial closure. The homogeneous appearance of cortex is probably secondary. PLATE 35 BULL. AMER. PALEONT., VOL. 69 e io" GRRE PEO Ry ties. © ben Sth: bet) Dae ‘ ~~ x s PLATE 36 BULL. AMER. PALEONT., VOL. 69 Figure ue CERIopoRID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 36 Leiosoecia parvicella (Gabb and Horn), 1860 ................000.. Lectotype, ANSP 31261, was probably figured by Gabb and Horn, 1861, pl. 69, figs. 36-38, from the ?Paleocene, Vincentown formation, Timber Creek, New Jersey. lias He: if lg. th. Zoarial fragment 2 showing the intersection and anasto- mosis of distally growing branches. Surface of zoarium <5. Surface of zoarium X30. Transverse thin-section 30. Zooecia in endozone have polygonal outlines; in exozone, zoecial growth slightly un- dulatory. Zooecial walls have submoniliform to moniliform profiles. Longitudinal thin-section X30. Zooecial walls thin and parallel-sided to submoniliform in endozone, thickened and having moniliform profiles in exozone. Note slight asym- metry of thickness across zooecial boundary zone. Thin inter- mediate diaphragms seen in most zooecia. Tangential thin-section 30. Interzooidal pores uncommon. View shows no apparent arrangement of large and small polymorphs. Longitudinal thin-section 200 showing microstructure, in- terzooidal pore, and thin intermediate diaphragms suboral to skeletal aperture. Tangential thin-section 100. Shows zooecial wall with amalgamate appearance and thin, dark zooecial linings. 205 206 Figure BULLETIN 291 EXPLANATION OF PLATE 37 1. Parleiosoecia jacksonica Canu and Bassler, 1920 .............0..00.00.... Lectotype, USNM Loc. 2933B-1, previously figured by Canu and Bassler, 1920, pl. 148, fig. 2, from the Eocene, Jacksonian, Eutaw Springs, South Carolina. tae 1b. lc. 1d. ter ils 1g. Zoarium 5. Zoarium has a large encrusting base and a single upright branch. Zoarial surface X30. Apertures of large polymorphs com- monly have slightly raised rims. Each large polymorph is surrounded by numerous small polymorphs. Tangential thin-section 30. Note discontinuous variation in size between large and small polymorphs. Longitudinal thin-section at base of erect branch «30. Shows hollow axial chambers formed by distal extension of basal layer. Transverse thin-section 30. Shows hollow axial! chamber, thin-walled polygonal zooecia in endozone, and thicker sub- moniliform walls in exozone. Longitudinal thin-section 100. Shows cylindrical cortex structure composed of subgranular calcareous tissue. Tangential thin-section 100. Cortex of large polymorphs distinct in appearance from indistinctly laminated walls of small polymorphs. PLATE 37 BULL. AMER. PALEONT., VOL. 69 % “ BULL. AMER. PALEONT., VOL. 69 PLATE 38 Figure 1-4, CERIOPORID CycLostomeEs (Bryozoa): NYE EXPLANATION OF PLATE 38 Parleiosoecia jacksonica Canu and Bassler, 1920 ..........00..0000.00..... All specimens figured are topotypes, identified by R. S. Bassler, and collected from the Eocene, Jacksonian, Eutaw Springs, South Carolina. 1. 3b. 4a. 4b. 4c. USNM 2933A. Zoarial fragment 5 showing branching, and intersection and anastomosis of distally growing branches in upper right. USNM 2933A-9. Longitudinal section, acetate peel X5 show- ing intersection and anastomosis of distally growing branches in upper right. The growing tip of the right-hand vertical branch abuts endozonal zooecia in the upper hori- zontal branch. USNM Loc. 2933A-23. Longitudinal thin-section 30 showing intrazoarial over- growth. Note that zooecial chamber in lower portion of figure is continuous with a zooecial chamber in the over- growth; other encrusted zooecia are sealed orally by thin, intermediate (?) diaphragms. Longitudinal thin-section 100 from same branch as above showing profile and structure of zooecial walls. Note thin intermediate diaphragms in terminal position, and two intermediate diaphragms in zone of zooecial bend. Also note wall structure of large polymorph in upper part of figure showing distinct cortex structure, and the steeply dipping laminae in adjacent wall of small polymorph. USNM 2933A-26. Tangential section, acetate peel 300 showing wall structure of large polymorphs and small polymorphs. Small poly- morphs distinctly laminate; cortex of large polymorphs subgranular. Longitudinal thin-section X30 showing axial chambers formed by basal layer and the intersection and anasto- mosis of two separate branches. A dark line marks surface of intersection in central and inner peripheral area but is lost in outer peripheral area. Note nearly 180° reflection of zooecial growth axes in lower part of figure so that zooecia grow proximally in relation to growth of branches. Detail of 1b X100 showing wedging out of zooecial cavities at surface of intersection. Zooecial walls merge continuously at surface of intersection to form a thin common wall. 207 208 Figure 1-5. Parleiosoecia jacksonica Canu and Bassler, 1920 BULLETIN 291 EXPLANATION OF PLATE 39 Figs. 1, 3-5 are specimens from the Eocene, Jacksonian, Eutaw Springs, South Carolina; fig. 2, specimens from Eocene, Jack- sonian, Santee River, three miles above Lenuds Ferry, South Carolina. 1. USNM Loc. 2933A, branch X5. 2. USNM Cat. 65447-1, figured by Canu and Bassler, 1922, pl. 148, fig. 1. 2a. Surface of branch 30. Shows apertures of large polymorphs with slightly raised rims surrounded by apertures of small polymorphs. Note regular, nearly rhombohedral, arrange- ments of large polymorphs. 2b. Longitudinal thin-section 100 showing axial chambers formed by basal layer. Note dark line separating basal layer from recumbent zooecial wall, and intermediate diaphragms in zone of zooecial bend. 2c. Detail of 2b 300 showing laminated structure of basal layer. 3. USNM Loc. 2933A-14. Longitudinal section, acetate peel <5 showing characteristic appearance of branch axis and coaxial exozone-endozone. 4. USNM Loc. 2933A-26. Longitudinal thin-section 100 show- ing axial chambers formed by basal layer. Note distal flexure of basal layer in center. 5. USNM Loc. 2933A-23. Transverse thin-section 100 show- ing basal layer and shape and arrangement of zooecial openings. BuLL. AMER. PALEONT., VOL. 69 PLATE 39 BULL. AMER. PALEONT., VOL. 69 PLATE 40 Figure CERIOPORID CycLosTomMeEs (Bryozoa): NYE EXPLANATION OF PLATE 40 1. Parleiosoecia jacksonica Canu and Bassler, 1920 .............00000000.... USNM Cat. 65449-1, Eocene, middle Jacksonian, 18 miles west of la. 1b. Ic. 1d. le. ibe Wrightsville, Georgia. Specimen figured by Canu and Bassler, 1922, pl. 148, fig. 6. Branch with brood chamber 5. Detail of brood chamber in la X30. Roof is partially broken, showing intrachamber zooecia. Longitudinal thin-section of same brood chamber X30; sec- and intrachamber zooecia. Detail of 1c 300 showing structure of brood chamber. Detail of 1c 300 showing structure of brood chamber roof and intrachamber zooecia. Longitudinal thin-section 300 showing wall structure of small polymorphs and terminal diaphragms. Proximal cham- ber has two diaphragms. USNM Cat. 65446, Eocene, middle Jacksonian, Rich Hill, 5% miles southeast of Knoxville, Georgia. Specimen figured by Canu and Bassler, 1922, pl. 148, fig. 3. Surface of branch showing brood chamber X30; most of roof is gone. A single zooecium opens into the central portion of brood chamber floor, but is not seen in this view. 209 210 Figure BULLETIN 291 EXPLANATION OF PLATE 41 1. Reptonodicava globosa (Michelin), 1846 ....0000.0...0000.ctee MNHN d’Orb. Coll. 2988-1, Jurassic, Bathonien, Luc (Calvados), France. la. 1b. lic: 1d. le. if 1g. ih. Clavate zoarium 2.5. Surface is smooth. Surface of zoaria] fragment X30. Longitudinal thin-section <5. Faint, dark-colored bands symmetrical with zoarial surface are probably abandoned growth surfaces. Outermost living chambers are closed aborally by basal diaphragms, and chambers are filled with dark, fine-grained sediments. Interior zooecial cavities are filled by clear calcite. Tangential thin-section 30. Longitudinal thin-section 100 showing thin, dark, basal diaphragms; diaphragms flex orally and form abutments. Longitudinal thin-section, crossed nicols 400. Laminae ap- pear to be oriented orally acute from boundary zone, then arch broadly across the cortex and are directed aborally in the inner cortex. Longitudinal thin-section 30. Shows submoniliform to moniliform zooecial walls; repetition of dark bands formed by a thin, dark zone with zooecial wall (detail of 1c) sug- gests cyclic growth intervals. Transverse thin-section X30 showing moniliform profile of zooecial walls and basa! diaphragms. PLATE 41 BULL. AMER. PALEONT., VOL. 69 BuLL. AMER. PALEONT., VOL. 69 PLATE 42 Figure i, CeRIoporID CycLostoMEs (Bryozoa): NYE EXPLANATION OF PLATE 42 Reptonodicava globosa (Michelin), 1846 000.0000... MNHN d’Orb. Coll. 2988-2, Jurassic, Bathonien, Luc (Calvados), France. la. 1b. tic: 1d. le. 1f. 1g. Zoarium X2.5. Zoarium is a hemispherical mass raised slightly above the substrate on narrower peduncle-like base. Surface of zoarium X30. Longitudinal thin-section 5. Intrazoarial overgrowth is seen in the proximal portion of colony. Longitudinal thin-section X20. Zooecial walls have sub- moniliform to moniliform profiles, walls typically equal and symmetrical in thickness across the boundary zone. Tangential thin-section 30. Longitudinal thin-section 100. Shows essentially continu- ously growing zooecial wall on right, and zooecial walls budding from basal layer on left. Zooecial wall, growing continuously, shows little variation in thickness. Zooecial walls newly budded from basal layer are very thin initially, but attain normal exozonal thickness and appearance within a very short distance. Longitudinal thin-section 100. Zooecia grew towards upper right. Walls of encrusting zooecia are continuous with thin, basal layer. Terminal diaphragms of subjacent zooecia are separated from encrusting basal layer by a dark line. 241 212 Figure BULLETIN 291 EXPLANATION OF PLATE 43 1-3. Reptonodicava globosa (Michelin), 1846 220.000.0000. All specimens from Jurassic, Bathonien, Luc (Calvados), France. ie 3b. 3c: MNHN d’Orb. Coll. 2988-1. Longitudinal thin-section 400 showing basal diaphragm. Diaphragm flexes and extends orally as a thin abutment. Zooecial walls nearly same color as clear calcite filling zooecial chamber, but show greater density of small dark granules. MNHN d’Orb. Coll. 2988-2, tangential thin-section 100. Numerous, short, blunt, mural spines give slightly crenulate appearance to outline of zooecial cavity. Note fine lamina- tions in outer cortex and zooecial lining. MNHEN d’Orb. Coll. 2988-3. Longitudinal thin-section 5. Shows typical growth habit and local intrazoarial overgrowth in upper left portion of zoarium. Fine-grained, dark-colored sediment fills last- formed living chambers and portions of zooecial cavities just subjacent to local intrazoarial overgrowth. Longitudinal thin-section 30, detail of local intrazoarial encrustation shown in 3a. Outer zooecial chamber subjacent to overgrowth is closed aborally by intermediate diaphragms and filled with dark, fine-grained sediment. Longitudinal thin-section 100. Shows zooecial walls with moniliform profile and interzooidal pores. Thin, basal dia- phragms are poorly preserved. Zooecial walls show obscure lamination. Longitudinal thin-section «100. Zooecia grew towards upper right. Note diaphragm subjacent to basal layer of intra- zoarial encrustation. Diaphragm shows slight aboral flexure at juncture with zooecia] wall and is apparently continuous with it. PLATE 43 BuLL. AMER. PALEONT., VOL. 69 BuLL. AMER. PALEONT., VOL. 69 PLATE 44 ~ ; a ecg weg \ bode Sapient 7 «eS cr fi Figure CERIOPORID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 44 1,2. Reptonodicava globosa (Michelin), 1846 .0..0000000...cccceeeeeeeeeeee Both specimens from Jurassic, Bathonien, Ranville (Calvados), France. ie Ze 2a 2b. Ze; USNM 32171-4, longitudinal section 5. Section includes encrusting basal portion. USNM 32171-1. Longitudinal thin-section 5, branch from a more massive- appearing zoarium. Dark bands suggest periodic addition of growth increments. Note numerous, evenly spaced basal diaphragms and very long, relatively straight zooecial lo- cated near major vector of branch growth. Longitudinal thin-section 5; another branchlike extension from the same zoarium. Various stages in zoarial growth are shown by dark bands, indicating that once budded zooecia continued to grow for most of the life of the colony, but grew at different rates. This part of the zoarium is overgrown by an encrusting cyclostome, c.f. Berenicea. Longitudinal thin-section 20, detail of 2a. Appearance of banding separating growth increments is partly due to in- clusion of dark material in walls, gradual increase in wall thickness in a given increment continued orally by re- duced thickness, and regular spacing of diaphragms. Major increments apparently include many minor incremental episodes. 213 214 Figure BULLETIN 291 EXPLANATION OF PLATE 45 1. Tetrocycloecia dichotoma Canu,, 1917 ......4..)...::.493 ee NMW 1859 L686-1, misidentified by Reuss as Heteropora dicho- toma (Goldfuss), from Miocene, Tortonian, Leithakalke, Eisen- stadt, Hungary. la. 1b. lic 1d. le. if 1g. Branch 5 showing regular distribution of apertures of large polymorphs. Tangential thin-section <30 showing discontinuous variation in size between openings of large and small polymorphs. Surface of zoarium X30. Surface is slightly worn. Note light-colored rings of calcareous tissue around apertures, ap- parently reflecting microstructure of zooecial wall; e.g. com- pare to le. Transverse thin-section 30. Exozone and endozone broad- ly gradational; zooecial walls in exozone submoniliform and slightly undulatory. Tangential thin-section 100. Zooecial walls in this view have integrate appearance because of concentration of dark granules along boundary zone; walls also have very thin, dark, laminate zooecial linings. Longitudinal thin-section 30. Note relatively simple ap- pearance, Exozone and endozone are gradational with broad zooecial curve from endozone; zooecia typically intersect zoarial surface at less than 90°. Zooecial walls display sub- moniliform profiles in both endozone and exozone. Longitudinal thin-section 100, detail of 1f. Shows zooecial wall with indistinct, orally convex laminae in cortex, and thin, dark, zooecial lining. PLATE 45 BULL. AMER. PALEONT., VOL. 69 a SAN ih PLATE 46 BULL. AMER. PALEONT., VOL. 69 Seay OBESE BEN ° Se broke vt, wed Fe & 8 255" ev 8 L he (@ ar igs Bret se Por @, Cras: | i YA< a ke f Jay ’ , 5S S ~ Sy) *, ~ = | - + a > & om, #s ce? fe Pm» 6 ~ : R : PF ; te, , Fay ps ci amat 9 Dar . : OF S03 a uA vs ; j , : 5 ; tal de valy z . ; , at < ~ ¥ af « Af ar: 2 aie o F “fs 4 es oN 2S ‘ - z x > io ae PG * - +s 4 { ¥ STS Be S . - 2X a s — ae + * f. q pe! f x ime. ae om i + Saad by Figure Crerioporip CycLosToMEs (Bryozoa): Nye EXPLANATION OF PLATE 46 1-3. Tetrocycloecia dichotoma Canu, 1918 200000... ccccccccceeeeeeteees All specimens figured were misidentified by Reuss as Heteropora dichotoma (Goldfuss), and were from the Miocene, Tortonian, Leithekalke, Eisenstadt, Hungary. Il la. 1b. Ic 2. 2a. 2b. 35 id. NMW 1859 L686-2. Branch <5. Longitudinal thin-section X30. Zooecial walls submoniliform in both endozone and exozone. Tangential thin-section 30. Transverse thin-section 30. NMW 1867-1, previously figured by Manzoni, 1877, pl. 12, fig. 46 as Heteropora dichotoma (Goldfuss). Branch X5. Longitudinal thin-section 30. Zooecia grow oblique from major vector of branch growth. NMW 1859 L686-1 30. Shows unworn portion of zoarial surface in lower right, and apertures of large polymorphs with slightly raised prominent rims. 25 216 Figure BuLLeETIN 291 EXPLANATION OF PLATE 47 1. Zonopora spiralis (Goldfuss), 1826. ..............c22..:2.2e ee Lectotype UB 133. Specimen is probably the specimen figured by Goldfuss, 1826, pl. 11, figs. 2a, b, from Cretaceous, Maastrich- tian, St. Petersberg near Maastricht (Limburg), Netherlands. tae 1b. ie: 1d. le. ibe 1g. Branch <5 showing typical helical appearance of zoaria. The convexly curved zoarial surface is referred to as the zoarial salient; the concave surface as the zoarial embay- ment. Terminal diaphragms appear as a nearly continuous sheet over the upper portion of the embayment. Surface of branch X20, detail of 1a. Deep tangential thin-section 30. Tangential thin-section 100. Deep tangential section of large zooecia showing amalgamate appearance and _ thin, laminate, zooecial linings. Transverse thin-section 30. 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 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. PLATE 47 BULL. AMER. PALEONT., VOL. 69 ie e, E aes so° : er: ‘ * sigs Fu BOR, Se aie PLATE 48 BuLL. AMER. PALEONT., VOL. 69 CERIopoRID CycLosToMEs (Bryozoa): NYE 27 EXPLANATION OF PLATE 48 Figure Page 1. Zonopora spiralis (Goldfuss), 1826 USNM Loc. 2965A-25, Cretaceous, Maastrichtian, Maastricht (Limburg), Netherlands. la. Longitudinal section, acetate peel x5. Shows profile with zoarial salients and embayments resulting from _ helical growth about a major vector of distal growth. Large zooecia intersect surface of salient at nearly right angles; small zooeCcia intersect surface of embayment obliquely. Branch to lower left appears to be an intrazoarial overgrowth. 1b. Tangential section, acetate peel 30. Shallow tangential section of small zooecia, deep tangential section of large zooecia. 1c. Longitudinal section, acetate peel 30, detail from 1a. Zoo- ecial walls in endozone thin and parallel-sided. Exozonal zooecial walls thicker, sometimes submoniliform, and have characteristic lanceolate profile. 1d. Longitudinal section, acetate peel X100, detail of 1c showing typical wall structure with light-colored cortex tissue bound- ed by thick, dark-colored deposits of zooecial lining. Spine- like lateral projections of cortex tissue submerged by zoo- ecial lining, but sometimes project into zooecial cavity as mural spines. le. Longitudinal section, acetate peel X100 showing wall struc- ture of small zooecia and interzooidal pores. Note thick deposits of dark-colored, longitudinally ]aminated tissue. 1f. Tangential section, acetate peel 100. Section is just distal to le, showing shallow tangential view of small zooecia. Note amalgamate, granular appearance of zooecial walls and thick zooecial linings. 1g. Tangential section, acetate peel 100. Section is moderately deep, showing thin zooecial lining. Note mural spines and narrow, straight, interzooidal pore in lower left quadrant of figure. 218 Figure 1k, BULLETIN 291 EXPLANATION OF PLATE 49 Zonopora spiralis (Goldfuss), 1826 .............0.cccccceeccccccseeceeeseeeeseaeees USNM Loc. 2965A-7, Cretaceous, Maastrichtian, Maastricht (Limburg), Netherlands. la. 1b. ic 1d. Longitudinal section, acetate peel 30. Transverse section, acetate peel X30. Large zooecia on left, small zooecia on right. Axis of branch is shifted to right of center; note thick zooecial linings in exozone. Tangential section, acetate peel X100, detail of large zooecia in 1b. Note increased thickness of zooecial lining in exozone. 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 zoarial salient. Longitudinal section, acetate peel 100. 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 nar- row, 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 sub- merged by thick zooecial linings in zooecial walls in right- hand portion of figure. PLATE 49 BULL. AMER. PALEONT., VOL. 69 dg P< id BX ED ss da yi eo . Bp Se , oy Ye . ae gy Ke BULL. AMER. PALEONT., VOL. 69 PLATE 50 ao 2 en ray: ay” > ys Ss OP ome St Figure CeRIoporID CycLosToMEs (Bryozoa): NYE EXPLANATION OF PLATE 50 1-3; Zonopora spiralis (Goldfuss); 1826. ......c.cc....00..ccseccseecrcecdesosstevseceeneees All specimens from Cretaceous, Maastrichtian, Maastricht (Lim- burg), Netherlands. iln lar 1b. 2a. 2b. Ze: 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. Longitudinal section, acetate peel 100, detail of 1a. Terminal diaphragm is porous and similar in appearance to zooecial lining of subjacent zooecial walls. Adjacent dia- phragms are not continuous across terminus of zooecial wall. Light-colored, homogeneous tissue of basal layer is con- tinuous with cortex tissue of encrusting zooecia. USNM Loc. 2965A-23. Longitudinal section, acetate peel 30. 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 laterally. Longitudinal section, acetate peel 100. Tissue of terminal diaphragm is continuous with zooecial linings, but does not extend across light-colored cortex tissue at terminus of zooecial wall. Longitudinal section, acetate peel 100. Detail of 1a show- ing cuspate extension of apertural rim and lateral flexure of zooecial lining to form terminal diaphragm. USNM Loc. 2965A-26, longitudinal section, acetate peel X100. Zooecial walls of large zooecia in outer exozone. The thin zooecial lining and rodlike extension of cortex into the zooecial cavity as mural spines, suggest a younger onto- genetic stage than shown in Pl. 36, figs. 1d, e, and pl. 37, fig. le. 219 220 Figure 2a. 2b. Ze: BULLETIN 291 EXPLANATION OF PLATE 51 Recent cerioporid, USNM 6086-1, 30. Brood chamber with roof partially broken away to show interior, revealing intra- chamber zooecia connected by thin septate walls.................... Recent cerioporid, BM O’Donoghue Coll. 1963-2-6-1 pt. all SCO OM Softitissues sare stained sie eeee Tangential thin-section. Basal portion of tentacular crowns in three zooecia; brown bodies in smaller zooecia to left. Dark lines in skeletal wall probably are algal or fungal borings. Note dark staining nuclei of cellular tissue in interzooidal part indicated by arrow. Longitudinai thin-section. Shows tentacular crowns and vis- ceral 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 Buti. AMER. PALEONT., VOL. 69 ee ie ~ RSA <9 ye INDEX Note: Light face figures refer to page numbers. Bold face figures refer to plate numbers. A Acamptostega ............ 63 Acanthoceramo- MOTE WAM coe re cevissesbscere 30 anomalopora, GeriOpora Assc-.ccs0s0s-- 13, 87, 88, 93, 114 IDitaxiawees ee 20-22 53, 87, 88-95, 189-191 Heteropora .............. 88 POlytaxdal ee.scc0cec se: 88 B Berenicea”™ 2iksecc..-: 44 213 beyrichi, Pennipora .. 56, 61, 176- 178 Cc Calyptrostega .............. 64 canui, Multigalea ........ 122 Cavatiaig eects. ere 130 Cellarian . 2 ciccenen 97 Ceramoporella ............ 30 Cerlocaval .iavieeesees. 1222; 27,305 40-43, 61, 79, 81, 98, 108,139 Ceriopora | ......26cisp20 Hise Glee eae 27, 52; 56, 61, 87, 112, 113, 120, 139, 140 Cerioporina .................. 8, 40, 64, 70 conifera, Millepora .... 140, 145, 147 corrugata, Densipora.. 131 Corymbopora .............. 10, 12, 27, tos 6< Corymbosal ...c3..2.020s 12, 62 corymbosa, Ceriocava. 2... 1-6 23, 32, 41, 42, 43-52, 67, 99, 108, 170-175 Milleporal .......2:....< 12, 40, 42, 43 Coscinoecia .................. 222272 305 38, 40, 43, 67, 79 crassa, Heteropora .... 60,61, 176- 178 cryptopora, Cerioporay Sees ie PPA SP Heteropora ...... 32-35 24, 53, 54, 56, 67, 114, 115- 122, 201-204 Defrancia 63, 64 Dendroecia Be coat ee uashoer eh 13, 41, 96, 98, 112 dichotoma, Cerioporal’ Vessels. 53, 114, 148, 150, 154 Heteropora .............. 113, 147, 148, 150, 214, 215 Tetro- cycloecia ........ 45,46 14, 147, 148- 155, 214, 215 Diplocava wy: eww aise. 18} OEY PAL Oe 80-81, 97, 140 Ditaxdaw ante 13. 14 22s 87-88, 113, 114 Domopora 2224s. 63, 64 dujardini, Fungella .... 63 Dysnoetopora’ .............. 10, 13 F falax, Ceriopora .......... 122 Fasciculipora .............. 62-64 frustulosa, Monticulipora ........ 42 Miumgeliian sok cecceuses 63, 64 fungiformis, Reptomulticavea .... 122, G globosa, Cerioporay: 5. 2-s.-: 14, 53, 55, 139-141, 145 Corymbopora .......... 17 Diplocava ................ 122 Reptonodicava 41-44 17,55, 140- 147, 210-213 grandis, Brachysoecia .......... 33 Grammascoecia .......... 53, 154 H Haploeeial sy csssccsoe LOMAS 22% Pp BYR obE Cal 81, 96-98, 112 Heteropora .................. 14, 21, 28, 52- 55, 61, 79, 87, 12-5. 1205 147 Heteroporina .............. 64 Heterotrypa ................ 114 | incondita, Diplocava ........ 16-19 13, 81-27, 122, 185-188 221 INDEX J jacksonica, Parleiosoecia ..37-40 14, 50, 67, 129-132, 135- 138, 206-209 E lamellosa, Multicrescis ................ 122 Meiosoeciaw 14, 21, 28, 70, 122-123, 129 M Macropora, Spiropora 131 magna, Heteropora .... 113 mamillosa, Reptonodicava ........ 139 mammulata, Monticulipora ........ 42 Meliceritities .............. 33 menardi, Corymbora ..... 10-12 12, 62, 64-70, 179-181 Corymbosa) 2.05... 64 Fasciculopora (Corymbosa) ............ 65 micropora, Ceriopora ........ 7-9 12, 53-62, 122, 140, 176- 178 Millenonaycn8 5022 oo. 41 Monticulipora ......... 41, 42 multilamellosa, Cerlocava’ ee. 13, 41, 96, 98, 106, 108, 112 Dendroecia .............. 106 Haploecia ....... 27-31 24, 50, 106- 112, 196-200 multitecta, Leptotrypella .......... 122 N neocomiencis, Corymbopora ..... 63, 64, 122 Radioporay occ... 122 nodosa, Seminodicrescis 131 Pp Pachystegay ee 64 pacifica, Heteropora . 113 Parleiosoecia ........... sa leale) Bale Ofey 79, 129-132 Partetrocycloecia ...... 148 parvicella, Heteropora) 123 Leiosoecia ........... 36 123-129, 205 Multicrescis 3... 14, 122, 123 pelliculata, Heteropora ce 113 Rolytaxiaeees sae 14, 87 pustulosa, Cavariaee. fe 131 Ceriopora, 2 42 Monticulipora .............. 42 R radiata Coscinoecia ....13-15 12, 23, 24, 25, 56, 70-79, 182-184 ramosa, Cavaria ......... 131 Reptomulticava .......... 61 Reptonodicava ............ 14, 23, 28, 30, 43, 52-55, 61, 81, 139-140 S Salplsinapes eee 97 Semilaterotubigera .... 130 spiralis Cerioporag 14, 155, 156 Spiroclatisa ..25)2.., Zonopora ........ 47-50 53, 156-162, 216-219 Spiroclausa,.7 4.) 47, 155 Stenolaemata .............. 19 straminea, Haploecia ........ 23-26 24, 26, 35, 46, 98-106, 192- 195 Milleporal, 20-2 13, 96, 98 Pustulopora ............ T Tabulipora <..63:..-c00c2.-: 46 Tetrocycloecia .......... 14, 28, 70, 123, 147-148, 150, 154 Tretocycloecia ............ 14, 147, 148 tuberosa, Multicrescis ............ 122 Tubuliporina .............. 63 Z ZONOPONAa she 14, 22, 28, 155 222 LI. LII. LITT. LIV. LV. LVI. LVII. LVITI. LIX. LX. LXI. LXIlI. LXIiII. LXIV. LXV. LXVI. LXVII. LXVIII. LXIX. (Nos. 231-232). BVONO) SOB; IO) PONS cece tts seen deere aeocece cer sce Antarctic bivalves, Bivalvia catalogue. (Nos. 233, 236). 387 Pp. 43 PIS. -.----------------ec-ccceseeeeereeceeecentees New Zealand forams, Stromatoporoidea, Indo-Pacific, Mio- cene-Pliocene California forams. (Nos. 237-238). CIS TB CYS) OSS, tereceessocpesenure oer eee Venezuela Bryozoa, Kinderhookian Brachiopods. (Nos. 239-245). ESO) ja)0b, SO) AWE ceeeeeceeceenoceee acer eee Dominican ostracodes, Lepidocyclina, mollusks. (Nos. 246-247). GIST, PAD SO ano Sve eee pe nace trees cteetet ccneemeceearenwons Cenozoic corals, Trinidad Neogene mollusks. (Nos. 248-254). Lip Pa iy sys yea) CRs parce en euetee ene eee neoprene Forams, North Carolina fossils, coral types, Cenozoic Echinoids, Cretaceous Radiolaria, Cymatiid gastropods (Nos. 255-256). Hailey ose Ga) 0) Stee eee ree sere eeeoee cece ooereeer Jurassic ammonites. (Nos. 257-262). B05) spies So wDISa aeeeeesee eee eoseestneesneseconsennseses Cretaceous Radiolaria and Forams, Pacific Silicoflagellates, North American Cystoidea, Cyclonema, Vasum. (No. 263). EINE S70) 015. peace eee cece cee etter Steere eta cere Bibliography of Cenozoic Echinoidea. (Nos. 264-267). BSS ID) OSie PS peeseese cas eae encnener ee cteene tame cateraron coe Radiolaria, cirripeds, Bryozoa, palynology. (Nos. 268-270). EACAGY eNO By ae TL 0) cies eee Oeen eee ee eke ee acon ree Mollusks, Murex catalogue, Cretaceous Radiolaria. (Nos. 271-274). 57/5 IDS Aang Sec eco ranee ne ee eee means Trace fossils, ammonoids, Silicoflagellates, microfauna. (Nos. 275-277). 320 pp., 56 pls. tafe d ete ee eossssesscennnssencernnnnsees Chitinozoa, Spumellariina, Mexican Ammonites (Nos. 278-281). 9 cesne-nesnesn-encensoseceresensceseessrennecrensenscnsunecnecnsaconsnesesaueess Palynology, corals, echinoderms, Foraminifera, and crinoids. (No. 282). CTIA 9) US dn) 0) IS id enero ee eee econ eee oct Ostracode Symposium. (Nos. 283-286). 639 Pp, 62, DUS. c2nscecenencescenenarcretecaranecerennossecceomwe Crinoids, gastropods, corals, ostracodes. (No. 287). 456 pps, 60. Pls. cocci nc cece mcerercteeeeneeeramenanawene Misc. Paleozoic (Nos. 288-290). OAM 0} RY An 0) (og beeen reece Skee eer Peon eee Paracrinoidea, ostracodes, cirripeds. (No. 291). DAIMID Sy RA la 2) (Sega ee cer cece eee ee ro PALAEONTOGRAPHICA AMERICANA Volume 1. See Johnson Reprint Corporation, 111 Fifth Ave., New York, Il. N. Y. 10003 Monographs of Arcas, Lutetia, rudistids and venerids. (Nos. 6-12). 5S Me PDS G md Le ee eee ce eeee eeremeecanaces Heliophyllum halli, Tertiary turrids, Neocene Spondyli, Paleozic cephalopods, Tertiary Fasciolarias and Pale- ozoic and Recent Hexactinellida. III. (Nos. 13-25). GTB, olby Ol OSs, ceececesecntececen beet eeeseer raccoons Paleozoic cephalopod structure and phylogeny, Paleozoic siphonophores, Busycon, Devonian fish studies, gastropod studies, Carboniferous crinoids, Cretaceous jellyfish, Platystrophia and Venericardia. IV. (Nos. 26-33). ENO) Soyo vst TIE) 0) ok een eer ee ee Rudist studies Busycon, Dalmanellidae Byssonychia, De- vonian lycopods, Ordovican eurypterids, Pliocene mol- lusks. V. (Nos. 34-47). EMTS oy ovata HO Ayo Kis eee peageemn eee ee ee ee Cee eee Tertiary Arcacea, Mississippian pelecypods, Ambonychiidae, Cretaceous Gulf Coastal forams. VI. (Nos. 38-41). 7 WW A oe | ol Kos ge ee pee eee eR PE Lycopsids and sphenopsids of Freeport Coal, Venericardia, Carboniferous crinoids, Trace fossils. VII. (Nos. 42-46). ZS TORT oN a} ts (At). 10) eee sso ee is eran ser Torreites Sanchezi, Cancellariid Radula, Ontogeny, sexual trilobites, Jamaician Rudists, crinoids. VIII. (Nos. 47, 48). 127M 26 0b ISe tree ees eek rece ee Gastropods, Devonian plants. 18.00 18.00 18.00 18.00 18.00 18.00 18.00 18.00 18.00 18.00 18.00 20.00 20.00 25.00 30.00 30.00 32.00 35.00 45.00 11.00 BULLETINS OF AMERICAN PALEONTOLOGY Vol. I-XXIII. See Kraus Reprint Corp., 16 East 46th St. New York, N. Y. 10017. U.S.A. XXIV. (Nos. 80-87). 334 ip phe (ap IS se ae eae ee te ee 12.00 Mainly Paleozoic faunas and Tertiary Mollusca. XOGV. | (Nos: 88-94B) 20) 306) ppryes Olsp isa ee er ees 12.00 Paleozoic, Mesozoic, and Miocene fossils. SOGVE.. ((Nos:295-100)5 4205 pp5 258 Nplssi ee -- 14.00 Florida Recent, Texas and South America Cretaceous, Cenozoic fossils. XOQVITS (Nos 101-108) gesii6iipps) 36" pls 14.00 Tertiary mollusks, Paleozoic Venezuela, Devonian fish. AXVIMI (Nos. 1095114). 412 pp,” 340 pis. sete ore 14.00 Paleozoic cephalopods, Cretaceous Eocene, forams. SOCEXG INOS} 1/5106) a V7 S Sepp amon DISae tere eee ... 18.00 Bowden forams and Ordovician cephalopods. EXOKOK: rag INOS RRUE 7 ren 65m DD sy O50 Ss, secrete eee eee ee 16.00 Jackson Eocene Mollusks. NKENONT./..\(NosasliiS-128) se 458h pps B27 uplsi gee ccace eee ee 16.00 Mollusks, crinoids, corals, forams, Cuban localities. OO coy (INOS5) 11292133) 5 02945 pp 7159s pls: pears ee eee 16.00 Silurian cephalopods, crinoids, Tertiary forams, Mytilarca. NEXEKTT Se (Nos: 34-139), 1443 spp:, Sl a pls age 16.00 Devonian annelids, Tertiary mollusks, Ecuadoran strati- graphy paleontology. XEXERTV ES (INOS: 1402145) 2 £4 00 ppml Sepia. ip tee cea. ee cee cee ocean reece 16.00 Forams, cephalopods, ostracods, conularid bibliography. OKO oe (Nos 6146-154) 5 e386 pp); 31 gpl ss cate ese eeee e - 16.00 Forams, cephalopods, mollusks, ostracods. XEXKENOVITS (Noss, 155-160) 24502) pps 53) PS gree tees: sees ceee ae ener caeeres eee 16.00 Forams, Eocene fish, rudists. XSCVINS (Noss 1612164); . 486-pp.,, 37 (pla: shoo ee 16.00 Cretaceous rudists, Foraminifera, Stromatoporoidea. DONC VIII: (Noss) 1652176)... A447 spp. e5:5i DS aiececcececnceeesee cee ee 18.00 Forams, ostracods, mollusks, Carriacou, fossil plants. SKEMOMIENS, (INosi9 177-183) 7 448 pps S6Upls se 2 ee ee 16.00 South American forams, Panama Caribbean mollusks. MU oCNo sel 84).7/7996 ‘pps daly oo a ce ee ee ene 18.00 Type and Figured Specimens P.R.I. DT (Nossal 85-192) oes Site pe e510 apse eee eee aes se 16.00 Forams, mollusks, carpoids, Corry Sandstone. DET SQNOS193)) oa. 4673 ppar4Saplett ee. ee ea eae ee eee eee 18.00 Venezuelan Cenozoic gastropods. SOIT © 1((Nos.9194-198) 0427p psn 29) pls) eet ee ee 16.00 Ordovician stromatoporoids, Indo-Pacific camerinids, Mis- sissippian forams, Cuban rudists. ALLY . oliNoss 199-203 565 spp. (68 ypl ss, aoe eee ees 16.00 Puerto Rican, Antarctic, New Zealand forams, Lepidocy- clina, Eumalacostraca. RL Ven (No: 9204) G4 pp. o6Seipls; evens n ee 2s 0) cere ee eee 18.00 Venezuela Cenozoic pelecypods. ALN. (Noss 205-281) 2419. ppv 70plss) a ee ee ee 16.00 Forams, Crustacea, brachipods, Recent mollusks. XLVI. (Noss 212-207)" S84" ppt, 33 spl st eee ee oe ee ee 18.00 Forams, mollusks, polychaetes, ammonites. AE VINT:: |: CNosa218)- 2 S05 eM ppt Styl sic ee eee 18.00 Catalogue of the Paleocene and Eocene Mollusca of the Southern and Eastern United States. ALIX. (Noss 219-224) 5 e674 top Soi pls eee ere ee 18.00 Peneroplid and Australian forams, North American car- poids, South Dakota palynology, Venezuelan Miocene mol- luska, Voluta. L. (No. 225-230). SiS Uipp:, 42 plist) en cerees eae ee ea ae 18.00 Venezuela, Florida cirripeds, forams, Linnaean Olives, Camerina, Ordovician conodonts. 4 - iM ria y) if i | a po i'B ny yest i na aE Py fe, > paeon _~ av \ i ny AY ‘Bil, in Ai my re ag ja yi Oh et u baler) me } ; ¥ v : - F wy a 7 vi Ne om Mi i) : cn ri 1 ri D 7 ‘ef Wi ' a ve “< Heme Bookbinding Co., Inc. 360 Summer Street Boston, Mass. 02210 r ANIL 3 2044 072 2 | | | | PPE NE, yh ees amie ne ETS IH SORE re oa SP ar hRehinll SKB Both Hy a Bian Teal tat Mt