TREATISE ON INVERTEBRATE pALEONTOLOGY
Part L
MOLLUSCA 4 Revised
Volume 4: Cretaceous Ammonoidea C. W. WRIGHT with J. ...
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TREATISE ON INVERTEBRATE pALEONTOLOGY
Part L
MOLLUSCA 4 Revised
Volume 4: Cretaceous Ammonoidea C. W. WRIGHT with J. H. CALLOMAN and M. K. HoWARTH
Prepared under Sponsorship of The Geological Society ofAmerica, Inc. SEPM (The Sedimentological Society) The Palaeontological Association
The Paleontological Society The Palaeontographical Society RAYMOND C. M ooRE
RoGER L. KAEsLER
Founder
Editor
ELIZABETH BROSIUS, }ILL HARDESTY, ]ACK KEIM }ANE KERNS, }ILL KREss, KAREN RENTERIA
Assistant Editors and Editorial Staff
THE GEOLOGICAL SOCIETY OF AMERICA , INC. and THE UNIVERSITY OF KANSAS BoULDER, COLORADO, and LAWRENCE, KANSAS
1996
© 1996 BY THE UNIVERSITY OF KANSAS AND THE GEOLOGICAL SOCIETY OF AMERICA, INC.
ALL RIGHTS RESERVED
Library of Congress Catalogue Card Number 53-12913 ISBN 0-8137-3112-7
Distributed by the Geological Society of America, Inc., P.O. Box 9140, Boulder, Colorado 80301, from which current price lists of parts in print may be obtained and to which all orders and related correspondence should be directed. Editorial office of the Treatise: Paleontological Institute, 121 Lindley Hall, the University of Kansas, Lawrence, Kansas 66045.
II
The Treatise on Invertebrate Paleontology has been made possible by ( 1 ) funding principally from the National Science Foundation of the United States in its early stages, from the Geological Society of America through the bequest of Richard Alexander Fullerton Penrose, Jr., and from The Kansas University Endowment Association through the bequest of Raymond C. and Lillian B . Moore; (2) contribution of the knowledge and labor of specialists throughout the world, working in cooperation under sponsorship of the Geological Society of America, the Paleontological Society, the SEPM (The Sedimentological Society), the Palaeontographical Society, and the Palaeontological Association; (3) acceptance by the University of Kansas of publication without any financial gain to the University; and (4) generous contributions by our individual and corporate sponsors.
Ill
TREATISE ON INVERTEBRATE PALEONTOLOGY Parts of the Treatise are distinguished by assigned letters with a view to indicating their systematic sequence while allowing publication of units in whatever order each is made ready for the press . Copies can be obtained from the Publication Sales Department, T he Geologi cal Society of America, 3300 Penrose Place, P. O. Box 9 1 40, Boulder, Colorado 8030 1. VOLUMES ALREADY P UBLISHED Part A. IKTRODUCTION, xxiii + 569 p., 37 1 fig., 1 979. Part C. PROTISTA 2 (Sarcodina, Chiefly "T hecamoebians" and Foraminiferida), xxxi + 900 p., 5,3 1 1 fig., 1 964. Part D. PROTISTA 3 (Chiefly Radiolaria, T intinnina), xii + 1 95 p., 1,050 fig., 1 9 54. Part E . ARCHAEOCYATHA, PORIFERA, xviii + 1 22 p., 728 fig., 1 9 5 5. Part E, Revised. ARCHAEOCYATHA, Volume 1, xxx + 1 5 8 p., 87 1 fig., 1 972. Part F. CoELENTERATA, xvii + 498 p., 2,700 fig., 1 956. Part F. COELENTERATA, Supplement 1 (Rugosa and Tabulata), xi + 762 p., 3,3 1 7 fig., 198 1 . Part G . BRYOZOA, xii + 2 5 3 p., 2,000 fig., 1 95 3 . Part G , Revised. BRYOZOA, Volume 1 (Introduction, Order Cystoporata, Order Cryptostomata), xxvi + 626 p., 1,595 fig., 1 983. Part H. BRACHIO PODA, xxxii + 927 p., 5, 1 98 fig., 1 96 5 . Part I . MOLLUSCA 1 (Mollusca General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda General Features, Archaeogastropoda, Mainly Paleozoic Caenogastropoda and Opisthobranchia), xxiii + 3 5 1 p., 1,732 fig., 1 960. Part K. MOLLUSCA 3 (Cephalopoda General Features, Endoceratoidea, Actinoceratoidea, Nautiloidea, Bactritoidea), xxviii + 5 1 9 p., 2,382 fig., 1 964. Part L. MOLLUSCA 4 (Ammonoidea), xxii + 490 p., 3,800 fig., 1 957. Part N. MOLLUSCA 6 ( Bivalvia), Volumes 1 and 2 (of 3), xxxvii + 952 p., 6, 1 98 fig., 1 969; Volume 3, iv + 272 p., 742 fig., 1 97 1 . Part 0. ART HROPODA 1 (Arthropoda General Features, Protarthropoda, Euarthropoda General Features, Trilobitomorpha), xix + 5 60 p., 2,880 fig., 1 9 5 9 . Part P. ARTHROPODA 2 (Chelicerata, Pycnogonida, Palaeoisopus), xvii + 1 8 1 p ., 565 fig., 1 95 5 . Part Q. ART HROPODA 3 (Crustacea, Ostracoda), xxiii + 442 p., 3,476 fig., 1 96 1 . Part R. ARTHROPODA 4, Volumes 1 and 2 (Crustacea Exclusive of Os tracoda, Myriapoda, Hexapoda), xxxvi + 651 p., 1 ,762 fig., 1 969. Part R. ARTHROPODA 4, Volumes 3 and 4 (Hexapoda), xxii + 6 5 5 p., 1 ,489 fig., 1 992. Part S. EcHINODERMATA 1 (Echinodermata General Features, Homalozoa, Crinozoa, exclusive of Crinoidea), xxx + 650 p., 2,868 fig., 1 967 [ 1 968]. Part T. ECHINODERMATA 2 (Crinoidea), Volumes 1 -3, xxxviii + 1 ,027 p., 4,833 fig., 1 978. Part U. EcHINODERMATA 3 (Asterozoans, Echinozoans), xxx + 695 p., 3,4 8 5 fig., 1966. Part V. GRAPTOLITHINA, xvii + 1 01 p., 358 fig., 1 95 5 . Part V, Revised. GRAPTOLIT HINA, xxxii + 1 63 p., 507 fig., 1 970. Part W MISCELLANEA (Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils, Problematica), xxv + 2 5 9 p., 1,058 fig., 1962. Part W, Revised. MISCELLANEA, Supplement 1 (Trace Fossils and Problematica), xxi + 269 p., 9 1 2 fig., 1 975. Part W, Revised. MISCELLANEA, Supplement 2 (Conodonta), xxviii + 202 p., frontis., 8 5 8 fig., 1 98 1 . IV
THIS VOLUME Part L, Revised. MoLLUSCA 4, Volume 4 (Cretaceous Ammonoidea), xx illus. on 2 1 6 fig., 1 9 96.
+
362 p., 2,070
VOLUMES IN PREPARATION Part B. PROTISTA 1 (Chrysomonadida, Coccolithophorida, Charophyta, Diatomacea, etc.). Part E, Revised. PORIFERA. Volume 2. Part G, Revised. BRYOZOA (additional volumes). Part H, Revised. BRACHIOPODA. Part I. Introduction to MoLLUSCA (part). Part J. MOLLUSCA 2 (Caenogastropoda, Srreptoneura exclusive of Archaeogastropoda, Euthyneura). Part L, Revised. MOLLUSCA 4 (Ammonoidea) (additional volumes) . Part M . MOLLUSCA 5 (Coleoidea). Part 0, Revised. ARTHROPODA 1 (Trilobita). Part Q, Revised. ARTHROPODA 3 (Ostracoda) .
EDITORIAL PREFACE Part L. Other volumes planned for the series are an introductory volume, volume 2 on Paleozoic ammonoids, and volume 3 on Tri assic and Jurassic ammonoids. In a way, it is unfortunate that the fourth volume i n the series is being published first, but even more unfortunate would be to delay publication of Dr. Wright's manuscript, which brings infor mation on the ammonoids up to date, a group of fossils that is vitally important for stratigraphy.
From the outset the aim of the Treatise on Invertebrate Paleontology has been to present a comprehensive and authoritative yet com pact s tatement of k nowledge concerning groups of invertebrate fossi ls . Typically, preparation of early Treatise volumes was undertaken by a single specialist with a syn optic view of the group being monographed. More rarely, two or perhaps three specialists worked together. Recently, however, both new Treatise volumes and revisions of exist ing ones have been undertaken increasingly by teams of specialists led by a coordinating author. Part L, Mollusca 4(4) , Revised has been prepared by a single author, Dr. C. W. Wright, and even this volume is p art of a much larger project on all the ammonoids that involves a number of specialists . Nev ertheless, few paleontologists have such an all-encomp assi n g command of a m aj o r group o f fossils as Dr. Wright has of the Cre taceous ammonoids. We are indeed privi leged that he has found both the time and the energy over the years to compile this in formation and share it with the paleontologi cal and geological communities. This volume on the Cretaceous Ammo noidea is the final section of the revision of
ZOOLOGICAL NAMES Questions about the proper use of zoo logical names arise continually, especially questions regarding both the acceptability of names and alterations of names that are al lowed or even required. Regulations pre pared by the International Commission on Zoological Nomenclature (ICZN) and pub lished in 1 98 5 in the International Code of Zoological Nomenclature, hereinafter referred to as the Code, provide p rocedures for an swering such questions. The prime objective of the Code is to promote stability and uni versality in the use of the scientific names of animals, ens uri ng als o that each generic name is distinct and unique, while avoiding v
unwarranted restrict ions on freedom of thought and action of systematists. Priority of names is a basic principle of the Code, but under specified conditions and by following prescribed procedures, priority may be set aside by the Commission. These procedures apply especially where slavish adherence to the principle of priority would hamper or even disrupt zoological nomenclature and the information it conveys. The Comm ission, ever aware of the changing needs of systematists, is undertak ing a revision of the Code that will further enhance nomenclatorial stability. Neverthe less, the nomenclatorial tasks that confront zoological taxonomists are formidable and have often justified the complaint that the study of zoology and paleontology is too of ten merely the study of names rather than the study of animals. It is incumbent upon all systematists, therefore, at the outset of their work to p ay careful attention to the Code to enhance stability by minimizing the number of subsequent changes of names, too many of which are necessitated by insuffi cient attention to detail. To that end, several pages here are devoted to aspects of zoologi cal nomenclature that are j udged to have chief importance in relation to procedures adopted in the Treatise, especially in this vol ume. Terminology is explained, and ex amples are given of the style employed in the nomenclatorial parts of the systematic de scriptions.
tam is the provision that within each group the categories are coordinate, that is, equal in rank, whereas categories of different groups are not coordinate. FORMS OF NAMES
All zoological names can be considered on the basis of their spelling. The first form of a name to be p ublished is defi ned as the original spelling ( Code, Article 3 2), and any form of the same name that is published later and is different from the original spelling is designated a subsequent spelling ( Code, Ar ticle 33). Not all original spellings are cor rect, nor are all subsequent spellings. Original Spellings
If the first form of a name to be published is consistent and unambiguous, the original is defined as correct unless it contravenes some stipulation of the Code (Articles 1 1 , 27 to 3 1, and 34) or unless the original publica tion contains clear evidence of an inadvert ent error in the sense of the Code, or, among names belonging to the family-group, unless correction of the termination or the stem of the type genus is required. An original spell ing that fails to meet these requirements is defined as incorrect. If a name is spelled in more than one way in the original publication, the form adopted by the first reviser is accepted as the correct original spelling, provided that it complies with mandatory stip ulations of the Code (Articles 1 1 and 24 to 34). Incorrect original spellings are any that fail to satisfy requirements of the Code, represent an inadvertent error, or are one of multiple original spellings not adopted by a first re viser. These have no separate status in zoo logical nomenclature and, therefore, cannot enter into homonymy or be used as replace ment names; and they call for correction. For example, a name originally published with a diacritical mark, apostrophe, diaeresis, or hy phen requires correction by deleting such features and uniting parts of the name origi nally separated by them, except that deletion of an umlaut from a vowel in a name derived
GROUPS OF TAXONOMIC CATEGORIES
Each taxon belongs to a category in the Linnaean hierarchical classification. The Code recognizes three groups of categories, a species-group, a genus-group, and a family group. Taxa oflower rank than subspecies are excluded from the rules of zoological no menclature, and those of higher rank than superfamily are not regulated by the Code. It is both natural and convenient to discuss nomenclatorial matters in general terms first and then to consider each of these three, rec ognized groups separately. Especially imporVI
from a German word or personal name un fortunately requires the insertion of e after the vowel. Where original spelling is j udged ro be incorrect solely because of inadequacies of the Greek or Latin scholarship of the au thor, nomenclatorial changes conflict with the primary purpose of wological nomencla ture as an information retrieval system. One looks forward with hope to a revised Code wherein the rules enhance stability rather than classical scholarship, thereby facilitating access to information.
AVAILABLE AND UNAVAILABLE NAMES
Editorial prefaces of some p revious vol umes of the Treatise have discussed in appre ciable detail the availability of the m any kinds of zoological names that have been proposed under a variety of circumstances. Much of that information, while important, does not pertain to the present volume, in which the author has used fewer terms for such names. The reader is referred to Part G, Bryozoa (Revised) of the Treatise and to the Code (Articles 1 0 to 20) for further details on availability of names. Here, suffice it to say that an available zoological name is any that conforms to all mandatory provisions of the Code. All zoological names that fail to com ply with mandatory provisions of the Code are unavailable and have no status in zoologi cal nomenclature. Both available and un available names are classifiable into groups that have been recognized in previous vol umes of the Treatise, although not explicitly differentiated in the Code. Among names that are available, these groups include invio late names, perfect names, imperfect names, vain names, transferred names, improved or corrected names, substitute names, and con served names. Kinds of unavailable names include naked names (see nomina nuda be low), denied names, impermissible names, null names, and forgotten names. Nomina nuda include all names that fail to satisfy provisions stipulated in Article 1 1 of the Code, which states general requirements of availability. In addition, they include names published before 19 3 1 that were un accompanied by a description, definition, or indication ( Code, Articles 1 2 and 1 6) and names published after 1 930 that ( 1 ) lacked an accompanying statement of characters that differentiate the taxon, (2) were without a definite bibliographic reference to such a statement, (3) were not proposed expressly as a replacement ( nomen novum) of a preexist ing available name ( Code, Article 1 3a), or (4) for genus-group names, were unaccompa nied by definite fixation of a type species by original designation or i ndication ( Code,
Subsequent Spellings
If a subsequent spelling differs from an original spelling in any way, even by the omission, addition, or alteration of a single letter, the subsequent spelling must be de fined as a different name. Exceptions include such changes as an altered termination of adjectival specific names to agree in gender with associated generic names; changes of family-group names to denote assigned taxo nomic rank; and corrections that eliminate originally used diacritical marks, hyphens, and the like. Such changes are not regarded as spelling changes conceived ro produce a different name. In some instances, however, species-group names having variable spell ings are regarded as homonyms as specified in the Code (Article 5 8) . Altered subsequent spellings other than the exceptions noted may be either inten tional or unintentional. If "demonstrably intentional" ( Code, Article 33, p. 73), the change is designated as an emendation. Emendations may be either j ustifiable or unjustifiable. Justifiable emendations are corrections of incorrect original spellings, and these take the authorship and date of the original spellings. Unjustifiable emendations are available names with their own author and date of publication. They are j unior ob jective synonyms of the name in its original form. Unintentional, subsequent, incorrect spellings are not available. They do not en ter into homonymy and cannot be used as replacement names. VII
Article 1 3b) . Nomina nuda have no status in nomenclature, a nd they are not correctable to establish original authorship and date. VALID AND INVALID NAMES
lm portan t cons iderati ons disti nguish valid from available names on the one hand and invalid from unavailable names on the other. Whereas determination of availability is based entirely on objective considerations guided by articles of the Code, conclusions as to validity of zoological names may be partly subjective. A valid name is the correct one for a given taxon, which may have two or more available names but only a single cor rect, hence valid, name, which is also gener ally the oldest name that it has been given. Obvious ly, no valid name can als o be an unavailable name, but invalid names may be either available or unavailable. It follows that any name for a given taxon other than the valid name, whether available or unavailable, is an invalid name. One encounters a sort of nomenclatorial no-man's land in considering the status of such zoo logical names as nom ina dubia (doubtful names), which may include both available and unavailable names. The un available ones can well be ignored, but names considered to be available contribute to un certainty and instability in the systematic lit erature. These can ordinarily be removed only by appeal to the ICZN for special ac tion. Because few systematists care to seek such remedy, s uch i nvalid but available names persist in the literature. A few such names are found in this volume. NAME CHANGES IN RELATION TO GROUPS OF TAXONOMIC CATEGORIES
the fo rm of adj ectival sp ecific names is changed to agree with the gender of a generic name in transferring a species from one ge nus to another, one need never label the changed name as nomen correctum. Similarly, transliteration of a letter accompanied by a diacritical mark in the manner now called for by the Code, as in changing origi n al ly broggeri to broeggeri, or eliminating a hy phen, as in changing originally p ublished cornu-oryx to cornuoryx, does not require the designation nomen correctum. Of course, in this age of computers and electronic data bases, such changes of name, which are per fectly valid for the purposes of scholarship, run counter to the requirements of nomen clatorial stability upon which the prepara tion of massive, electronic data bases is predi cated. Genus-Group Names
Conditi ons warranti n g change of the originally published, valid form of generic and subgeneric names are sufficiently rare that lengthy discussion is unnecessary. Only elimination of diacritical marks and hyphens in some names in this category and replace ment of homonyms seem to furnish basis for valid emendation. Many names that for merly were regarded as homonyms are no longer so regarded, because two names that differ only by a single letter or in original publication by the presence of a diacritical mark in one are now construed to be entirely distinct. As has been pointed out above, difficulty typically arises when one tries to decide whether a change of spelling of a name by a subsequent author was intentional or unin tentional, and the decision has often to be � ade arbitrarily.
Species-Group Names Family-group Names: Authorship and Date
Detailed consideration of valid emenda tion of specific and subspecific names is un necessary here, both because the topic is well understood and relatively inconsequential and because the Treatise deals with genus group names and higher categories. When
All family-group taxa having names based on the same type genus are attributed to the author who first published the name of any of these groups, whether tribe, subfamily, or Vlll
the information of interest is the author who initially introduced a taxon rather than the author of the superfamily as defined by the Code. The latter is merely the individual who fi r st defi n ed some lower-ranked, family group taxon that contains the nominotypical genus of the superfamily. On the other hand, the publication that introduces the super family by nomen translatum is likely to fur nish the information on taxonomic consid erations that support definition of the taxon. An example of the use of nomen translatum is the following.
�-amily (superfamily being almost inevitably later-conceived taxon) . Accordingly, if a iamily is divided into subfamilies or a sub t-amily into tribes, the name of no such sub tamily or tribe can antedate the family name. .\[oreover, every family containing differen t iat ed s ubfam ilies m us t have a nom i no n·p ical subfamily (sensu stricto) , which is 6ased on the same type genus as the family. F inally, the author and date set down for the nominotypical subfamily invariably are iden tical with those of the family, irrespective of whether the author of the family or some erfamily ranging from the Barremian to the \ !aasrrichtian. They are described in full in :his volume. Ammonitina likewise range from the base :> r- the Jurassic to the top of the Cretaceous. A few survivors of the largely Jurassic Hap : oceraraceae, Haploceratidae and Oppeliidae Srreblitinae), range into the Cretaceous and J.re described in this volume with the wholly C retaceous Oppeliidae (Aconeceratinae) and Binneyitidae. The Perisphinctaceae are pre .iominately Jurassic, but three important :".imilies-Polyptychitidae, Olcostephanidae, .;.:1d :"eocomitidae-are mainly Lower Cre :.;.,:eous and, with minor derivative families,
are fully described here. Desmocerataceae, Pulchelliaceae, Hoplitaceae, and Acantho cerataceae are wholly Cretaceous and are dealt with in this volume. Ancyloceratina had a few late Jurassic members but flourished in the Cretaceous and are all described in this volume. Suborder PHYLLOCERATINA Arkell, 1 9 5 0 Superfamily PHYLLOCERATACEAE Zittel, 1 884 Family PHYLLOCERATIDAE Zittel, 1884 Subfamily PHYLLOCERATINAE Zittel, 1884 Phylloceras (Phylloceras) SUESS, 1 865 [*Ammonites heterophyllu s J . SOWERBY, 1 820] [ = Rhacoceras HYAIT, 1 867; Xeinophylloceras B ucKMAN, 1 92 1 in 1 909- 1 9 30; Heterophylloceras KOVACS, 1 9 3 9 ; Pseu dophylloceras BEZNOSOV, 1957] . Lower C reta ceous (Berriasian-Barremian). P. (Partschiceras) FuciNI, 1923 in 1923-1935 [*Am monites partschi HAUER, 1 854, non KursTEIN, 1 843] [ =Procliviceras F uc iNI, 1 92 3 in 1 923- 1 9 35; Macrophylloceras SPAI'H, 1927a; Proclivioceras Ro MAN, 1 9 3 8 ; Partschiphylloceras ROMAN, 1 9 3 8 ; Adobofoloceras ]OLY, 1 977] . Lower C retaceous (Ber riasian)-Upper Cretaceous (Maastrichtian). P. (Zetoceras) KovAcs, 1 939 [*Ammonites zetes OR BIGNY, 1 850a] [ = Lavizzaroceras KOVACS, 1 93 9 ] . Lower C retaceous (Berriasian- Valanginian). Phyllopachyceras SPATH, 1 925e [*Ammonites in fUndibu lu m ORBIGNY, 1 8 4 1 ] . Lower C retaceous (Bar remian)-Upper Cretaceous (Maastrichtian). Hypophylloceras (Hypophylloceras) SALFELD, 1 924 [* Phylloceras onoense STANTON, 1 895] [= Goreto phylloceras COLLIGNON , 1 94 8 ; Euphylloceras DRUSHCH!TS, 1 953b; Aphroditiceras MAHMOUD, 1 955] . Lower C retaceous (Valanginian)-Upper C re taceous (Cenomanian). H . (Neophylloceras) SHIMIZU, 1 9 34 [*Ammonites (Scaphites?) ramosus MEEK, 1 857] [ =Paraphylloceras SHIMIZU, 1 935a, non SALFELD, 1 9 1 9 ; Hyporbu lites BREISTROFFER, 1 947a; Epiphyl/oceras COLLIGNON, 1956] . Lower C retaceous (Aptian)-Upper C retaceous (Maastrichtian). Carinophylloceras KLINGER, WIEDMANN, & KENNEDY, 1 975 [* C. col/ignoni KLINGER, WIEDMANN, & KEN NEDY] . Lower C retaceous (Lower Albian).
2
Cephalopoda-Cretaceous Ammonoidea
Subfamily CALLIPHYLLOCERATINAE Spath, 1927 Calliphylloceras SPATH, 1927a [* Phylloceras disputabile ZITTEL, 1869] [=Neocalfiphylfoceras BESAIRIE, 1936; Capitanioceras KovAcs, 1 939] . Lower C retaceous (Berriasian-Albian). Holcophylloceras SPATH, 1 927a [ * Phylloceras mediterraneum NE UMAYR, 1 8 7 1 ] [ = Salfeldiella SPATH, 1 927a; Telegdiceras KovAcs, 1 942] . Lower C retaceous {Berriasian-Aptian). Ptychophylloceras SPATH, 1927 a [* Phylloceras ftddeni WAAGEN, 1 8 75 in 1 873-1 875] [= Tatroceras KOVACS, 1939; Neu mayriceras SORRENTINO, 1 942, non RoLLIER, 1 909; Tatrophylloceras BEZNOSOV, 1957]. Lower C retaceous (Berriasian-Aptian). Sowerbyceras PARONA & BONARELLI, ·1 895 [*Ammo nites tortisu lcatus ORBIGNY, 1 84 1] [=Martelliceras SORRENTINO, 1 9 42 , non SCHINDEWOLF, 1 925; Gyrophyllites WIEDMANN, 1963] . Lower Cretaceous (Berriasian-Albian).
Suborder LYTOCERATINA Hyatt, 1 889 Superfamily LYTOCERATACEAE Neumayr, 1 875b Family LYTOCERATIDAE Neumayr, 18 75b Subfamily LYTOCERATINAE Neumayr, 18 75b Lytoceras SuEss, 1 865 [*Ammonites jimbriatu s J. SoWERBY, 1 8 1 7a] [= Ophiceras SuEss, 1 865; Thysa noceras HYATT, 1 867; Thysanolytoceras BuCKMAN, 1 905; Trachylytoceras BUCKMAN, 1 9 1 3 in 1 9091 930; Fimbrilytoceras B UCKMAN, 1 9 1 8 in 1 9091 930; Kal!ilytoceras BucKMAN, 1921 in 1909-1930; Lobolytoceras B ucKMAN, 1 923 in 1 909- 1 930; C osmolytoceras SPATH, 1924; Crenilytoceras B ucK MAN, 1 926 in 1 909-1930; Orcholytoceras BucKMAN, 1926 in 1909- 1 930; Saturnoceras ANDERSON, 1938; Biasaloceras ORUSHCHITS, 1 953a; Dinolytoceras BEZNOSOV, 1 956; Valento/ytoceras BEZNOSOV, 1 956] . Lower C retaceous (Berriasian)-Upper C retaceous (C enomanian). Pterolytoceras S PATH, 1 927a [*Ammonites exoticus OPPEL, 1 863 in 1 862-1 863]. Lower Cretaceous (Ber riasian- Valanginian). Eulytoceras SPATH, 1927 a [*Ammonites inaequalicosta tus ORBIGNY, 1 840]. Lower C retaceous (Hauterivian Lower Aptian). Hemilytoceras S PATH , 1 927a [*Ammonites immanis OPPEL, 1 865] . ?Lower C retaceous. Metalytoce ras S PATH, 1 927 a [* Lytoceras triboleti UHLIG, 1902] . Lower C retaceous {Valanginian). Ammonoceratites BOWDITCH, 1 822 [*Ammon ites lama rcki BoWDITCH, 1 822] [=Ammonoceras LA MARCK, 1 822] . Lower C retaceous (Albian).
Argonauticeras ANDERSON, 1938 [* Lytoceras argonau tarum ANDERSON, 1 902] [ = Pseudotetrago nites ORUSHCHITS, 1 956] . Lower C retaceous (Upper Aptian-Middle Albian). Pictetia UHLIG, 1883 [*C rioceras astierianum ORBIGNY, 1 842] . Lower C retaceous (Upper Aptian-Middle Albian). Protetragonites HYATT, 1900 [*Ammonites quadriscul catus ORBIGNY, 1 8 4 1 ] [= Hemitetragonites SPATH, 1927a; Leptotetragon ites SPATH, 1927a]. Lower Cre taceous (Berriasian-Upper Albian). Carinolytoceras WIEDMANN, 1 962c [* C . carina tum WIEDMANN, 1 9 62c] . Lower C retaceous (Upper Aptian).
Superfamily TETRAGONITACEAE Hyatt, 1 900 [nom.
transl WIEDMANN,
1962a, p. 1 47, exTeuagonitidae HYATT, 1900, p. 568]
More involute than Lytocerataceae, with whorl section less circular and becoming a ngled in some forms; several auxiliary saddles developing in suture line; shell lirate, striate, smooth, or occasionally strongly ribbed, and constrictions usually present. Lower Cretaceous (Barremian)-Upper Creta ceous (Maastrichtian). Derived from Protetragonites (Lytocer atidae) in the Barremian. The two families, Gaudryceratidae and Tetragonitidae, are closely related in the Lower Cretaceous but more distinct i n the Upper Cretaceous. Gaudryceratidae retain the q uinq uelobate primary suture of the Lytocerataceae, but some Tetragonitidae develop a sexlobate pri mary suture. HENDERSON, 1 970; KENNEDY & HENDERSON, 1 9 92a; KULLMANN & WIED MANN, 1 970; MuRPHY, 1 967c; WIEDMANN, 1 962a, 1 962b; WRIGHT & KENNEDY, 1 9 84; WRIGHT & MATSUMOTO, 1 9 54. Family GAUDRYCERATIDAE Spath, 1927 [Gaudryceratidae SPATH, 1927a, p. 661 [=Gaudryceratinae WRIGHT, 1 9 57b, p. 200; Hypogaudryceratinae SHIMIZU, 1934, p. 67; Kossmatellinae BREISTROFFER, 1953b, p. 7 1 ; Venebritinae WIEDMANN, 1962a, p. 150]
Whorl section typically rounded or ellip tical, sometimes depressed; shell ornamented with lirae or fine rib lets, o ccasionally smooth, and with constrictions; suture with symmetrically bifid saddles and a single main saddle in internal suture, but with additional
Lytoceratina- Tetragonitaceae small saddles in some forms. KENNEDY & KLINGER, 1 97 9 b; KENNEDY & S UMMES BERGER, 1 9 8 6 . Lower Cretaceous (Barre mian)-Upper Cretaceous (Maastrichtian). Eogaudryceras SPAT H , 1 9 27a, p. 66 [*Ammonites nu midus CoQUAND, 1 880, p. 22; 00) . Moderately evolure; whorl section rounded ro rectangular; smooth or ornamented with fine radial lirae; con strictions present in some and may become strong. KENNEDY & KLINGER, 1 979 b. Lower C retaceou s (Barremian-Upper Albian): France, Spain, England, Bulgaria, Caucasus, Italy, Switzerland, Algeria, South Africa (Zululand), Madagascar, Egypt (Sinai) , California, Antarctica (Alexander Island). E. (Eogaudryceras). Whorl section initially trap ezoidal, becoming rounded and sometimes com pressed in aduhs; fine, sinuous lirae projected s trongly fo rwards over venter, bur mold is smooth; constrictions weak or absent, typically on inner whorls only; suture with symmetrically bifid saddles and large suspensive lobe. MURPHY, 1 967 c. Occurrence and distribution as for ge nus. FIG. 1 , 3a-d. *E. (E.) nu midu m (Co QUAND), Barremian; a, b, lecrorype (designated HowARTH , herein), Algeria (Djebel-Ouach, Constantine), X I (Sayn, 1 890, pl. I , fig. 3a,b); c,d, France (Hyeges, Alpes-de-Haute Provence) , X I (Murphy, 1967c, pl. 5, fig. 1-2). E. (Eoretragonites) BREISTROFFER, 1947b, p. 72 (*E. raspaili; OD]. Whorl section initially rather de pressed and more angular or rectangular than in E. (Eogau dryceras) at all growth stages; outer sur face of shell smooth or finely striate; constric tions moderate ro strong at all growth stages and projected gendy forwards over venter; suture line with asymmetrically bifid saddles and in cipient second lateral saddle in internal suture. Derived from E. (Eogau dryceras) in rhe Upper Aptian. Lower C retaceous (Upper Aptian-Middle Albian): France, Spain, Switzerland, Bulgaria, Caucasus, South Mrica (Zululand), Madagascar, Egypt (Sinai), California, Antarctica (Alexander Island). F I G . I , I a, b. *E. (Eotetragonites) raspaili B REISTROFFER, Upper Aptian, France (Hyeges, Alpes-de-Haute Provence); XI (Kilian, 1 9 1 3 , pl. 1 1 , fig. 3 ) . FIG. l , lc,d. E. (E.) wintu niu m (ANDERSON), Upper Aptian, north ern California; XI (Murphy, 1 967c, pl. 3, fig. 5-6). Anagaudryceras SHIMIZU, 1 9 34, p. 67 [*Ammonites sacya FoRBES, 1 846, p. 1 1 3 ; OD] [=Paragaudryceras SHIMIZU, 1 934, p. 67 (type, Gau dryceras limatu m YABE, 1 903, p. 34; 00) ; Mu rphyelfa MATSUMOTO in MATSUMOTO, MURAMOTO, & TAKAHASHI, 1972, p . 208 (type, Kossmatella (Mu rphyella) enigma; OD)]. Whorl section always rounded; early whorls circular or depressed; later whorls compressed and higher; mold smooth, bur shell with very fine, pror siradiare or sinuous lirae; periodic, collared con snictions on inner whorls becoming more numer ous on o uter whorls, where they form broad, --
--
--
3
flanened, foldlike ribs. Derived from E. (Eogaudry ceras) in the early Albian. KENNEDY & KLINGER, 1979b; WRIGHT & KENNEDY, 1984; HENDERSON & McNAMARA, 1 9 8 5 . Lower C retaceous (Middle Albian)-Upper C retaceou s (Maastrichtian): England, France, Germany, Austria, Romania, Algeria, Angola, South Africa (Zululand ) , Madagascar, southeastern India, Japan, Sakhalin, Kamchatka, Alaska, B ritish Columbia, California, New Zealand, Antarctica (Seymour Island). FIG. 1 ,4a-d. *A. sacya (FORBES), Upper Albian or Cen omanian, southeastern India (Verdachellum); a,b, holorype, BMNH C . 5 1 067, X 1 (new); c, d, bolo type of A m monites bu ddha FoRBES, BMNH C.22673, X0.7 (new) . Gaudryceras GROSSOUVRE, 1 894, p. 225 [*Ammonites mitis HAUER, 1 866, p. 305; SO BOULE, LEMOINE, & THEVEN I N , 1 906, p. 1 1 ( 3 1 ) ) [ = Epigau dryceras S H IMIZU, 1 934, p. 67 (type, Lytoceras st riatu m }IMBO, 1 894, p. 3 5 ( 1 8 1 ); OD); Hemigau dryceras SHIMIZU, 1934, p. 67 (type, Lytoceras (Gau dryceras) den manense WHITEAVES, 1 9 0 I , p . 3 1 ; 00); Neogau dryceras S H IMIZU, 1 93 4 , p . 67 ( type, Gau dryceras tenu iliratum YABE, 1 903, p . 19; 00); Pseu dogau dryceras S H I M IZU, 1 9 34, p. 67 (type, Gau dryceras tenuiliratu m var. infrequens YABE, 1 903, p. 28; 00)) . Inner whorls evolure, serpenticone, rounded or depressed; later whorls expanding more rapidly and becoming compressed; ornamented with lirae or wirelike, flexuous ribs, single o r branched, which may coarsen and bunch on rhe outer whorls; variable, foldlike ribs may also de velop on rhe outer whorls; periodic constrictions on inner mold marked by depressions and collars on rhe shell surface. Suture with wide, bifid saddles and lobes and a retracted suspensive lobe. Lirae and ribs coarser than in Anagau dryceras. KENNEDY & KLINGER, 1 979b. Lower C retaceous (Upper Albian) Upper Cretaceous (Maastrichtian): worldwide. G. (Gaudryceras). Large, wirh constrictions; devel oping foldlike ribs on body chamber. KENNEDY & SUMMESBERGER, 1 979. Occurrence and disni burion as for genus. FIG. 2,4a,b. * G. (G.) mite (HAUER), Coniacian, Gosau Beds, Ausnia (Strobl, Wolfgangsee) ; holotype, X0.7 (Kennedy & Summesberger, 1979, pl. 1 ) . G. ( Mesogaudryceras) SPATH, 1927a, p . 6 6 [*Am monites leptonema SHARPE, 1 85 5 . p. 32; 00) . Smaller, more involute and expanding more quickly than G. (Gau dryceras}; whorls more compressed from an early stage, sometimes de veloping trigonal whorl section, widest near um bilicus, and with narrowly arched venter; lirae distant at first, then fine, dense, and snongly p rojected on rhe venter; no constrictions. WRIGHT & KENNEDY, 1 984. Upper C retaceou s (Lower Cenomanian-Middle Cenomanian): En gland, France, Germany, Romania, Ukraine (Crimea), Turkmenistan (Koper-Dag), Green land. F I G . 2 ,3a, b. * G. (M.) lept onema (SHARPE), Lower or Middle Cenomanian, Lower Chalk, England (Venrnor, Isle of Wight); bolo type, GSM 7762, X I . --
--
--
4
Cephalopoda-Cretaceous Ammonoidea
1d
1c
2c Vertebrites
Eotetragonites
4b
4c Eogaudryceras
Anagaudryceras
FIG. I . Gaudryceratidae (p. 3-5) Verrebrites MARSHALL, 1926, p. 1 3 8 [• I; SCHWEILER) ; a, X3; b, X7 (both McNamara, 1985 KoJoceras R ICCARJ)!, AGUIRRE URRETA , & M EDI S.,. 1 9 87 , p. 1 6 7 ( * K. talenkanum; O D ] . Venter rounded ro tabulate with intermittent, feeble keel: with falcate striae to weak ribs. Suture with lobes L and U2 trifid to subbifid. Lower Cretaceous (Upp� Albian}: Argentina (Patagonia), Angola. ·
- Ammonitina-Haplocerataceae
15
,.,"- ...... ,/' '
'
:
', \
\
\ \
'
.
\ I
I
\
'
' '
I I
:
\
Sanmartinoceras
3c I I
Gyaloceras
.,0
n
... Sa
Sbu
Falciferella
Theganoceras
Ac oneceras
FIG. 9 . Oppeliidae (p. 14)
Family BINNEYITIDAE Reeside, 1927 (BinncyiE,
1927o. p. 4}
strengthened or visible only on shoulders. Sutures simplifYing with one or more lobes bifid, rending ro have long, narrow, simple or
Small, compressed, Rat-sided, with Rat or
entire elements with parallel sides; second
sulcate venter and weak ornament of sinuous
lateral saddle projecting beyond first. Nacre
ro falcate striae or riblets; ornament may be
ous shell with metallic luster as
in
16
Cephalopoda-Cretaceous Ammonoidea
I
2a
,
2b Johnsonites
Binneyites
n 3b
�
{?�
i
3c
Borissjakoceras
4c
2d
Tropitoides
FIG. I 0. Oppeliidae & Family Uncertain (p. 16-17)
Aconeceratinae. Binneyitidae are derived fro m
the Aconeceratinae by way of
Falciferella. Lower Cretaceous (Upper Albian) Upper Cretaceous (Lower Santonian). Borissjakoceras ARKHANGELSK¥, 1 9 1 6 , p. 5 5 [*B. mirabilis; OD]. Moderately evolute to rather invo lute; Rat-sided; venter bluntly trapezoidal to rounded; surface smooth except for traces of pror siradiate ribs on shoulders. Suture simple with deep, narrow, parallel-sided L and broad saddles; saddle E/L bifid or entire, L/U trifid. Lowu Crttauous (Upper Albian)-Upper Cretaceous (Lower Turonian): France, Angola, South Mrica (Zululand), northern Siberia, Turkestan, Bathurst Island, New Zealand, Alaska, British Columbia, USA (Western Interior), Colombia, Argentina, Greenland.--FIG. 1 0,3a-
c. • B. mirabilis, Lower Turonian, Turkestan; a,b, X I ; c, X3 (all Arkhangelsky, 1 9 16). Johnsonites CoBBAN, 1961, p. 743 (*]. sulcatus; OD]. Flat-sided, with Aat co feebly sulcate veneer; smooch or with weak. falcoid ribs. Suture with deep, narrow external lobe divided by long, narrow foliole. Uppu Cretaceous (Upper Cenomanian): Wyoming, Colo rado.--FIG. i0,2a-d. ·;. sulcatus, Wyoming; a,b, X 1 ; c,d, X4 (all Cobban, 1961). Binneyices REESIDE, 1 927a, p. 4 [•B. parkensis; M]. Very involute; venter Aac; shoulders sharper and ventrolateral ornament stronger chan in Borissjnkouras. Suture with long, narrow, parallel sided, bifid or entire first lateral saddle, simple and broadening L, and very simple other elements; aux iliaries tending to increase in number with time. Upper Cretaceous (Lowu Turonian-Lower San tonian): USA (Western lnccrior).--FIG. 10,1a-c.
Ammonitina-Haplocerataceae *B. parkensis, Coniacian, Wyoming; a,b, X2; c, X4 (Reeside, 1927a).
Family Uncertain Tropitoides SPATH, 1 925e, p. 1 02 [*Ammonites obesus STOI.ICZKA, 1 86 5 , p. 5 5 ; 00] . I n volute, high whorled, with slightly convex sides sloping towards rather narrow venter bearing prominent, slightly rounded keel; dose, rather flat, falcoid ribs ending in feeble ventrolateral clavi. [Suture with second lat eral saddle projecting beyond other elements and the minute umbilicus recall Aconeceratinae but other features and stratigraphical horizon render connection unlikely.] Upper Cretaceous (?Middle Turonian, Upper Turonian}: France, Israel, southern India. F IG. l 0 , 4a-c. * T. obesus (SToLICZKA), ?Middle Turonian, southern India; a,b, X0.75; c, X I (all Kossmat, 1 895-1 898). --
Superfamily PERISPHINCTACEAE Steinmann, 1 890 Family POLYPTYCHITIDAE Wedekind, 1918 [nom. correct. SPATtl,
1 924a, p. 87, pro Polyprichitidae WEDEKJND, 1 9 1 8 , p . 103]
With the widespread retreat of the seas from many marginal areas of the northern hemisphere at the end of the Jurassic, the descendents of the boreal Dorsoplanitinae diversified in separate faunal provinces or subprovinces covering a rapidly changing pattern of partially isolated, shallow epicon tinental seas. Their remains are now found in scattered outcrops of highly incomplete, of ten condensed successions with phosphatic concretions. Difficulty in correlation is such that each faunal p rovince retains its own scheme of zonation. Six such provinces may be distinguished: ( 1 ) Spilsby (eastern En gland-eastern Greenland) ; ( 2 ) Lower Saxony; (3) Russian Platform; (4) Siberian Arctic (i ncluding the B a rents shelf and Spitsbergen) ; (5) Canadian Arctic (Sverdrup basin, Yukon) ; (6) northern Pacific cordillera (northern California, Oregon, Washington, British Columbia) . It is often difficult to decide to what extent faunal differences re flect differences in age or provincialism, and condensation can make it hard to establish the variability of contemporaneous assem-
17
blages or biospecies. Classification has there fore led to phylogenetic schemes of improb able complexity. Morphol ogical ly, the range of forms found in the family is unusually wide and untypical of perisphinctids in general. It in cludes involute, compressed homeomorphs of some Macrocephalitinae or Aulacoste phanidae ( Tollia) ; sphaerocones or cadicones that may be hard to distinguish from Cado ceratinae or Pachyceratidae (Polyptychites) ; serpenticones, homeomorphs of Stephano ceratinae (Simbirskites) ; and even oxycones, homeomorphic with Oxynoticeratinae (Platylenticeras) . Sometimes the whole range from platy- to sphaerocones is found within the variability of what is almost certainly but a single biospecies (e.g., Thorsteinssonoceras ellesmerense J ELETZKY, 1 965). Interruption of ribbing on the venter to give a smooth band evolved several times in the family and has therefore as little systematic value here as elsewhere in the Perisphinctaceae. Constric tions similarly came and went. The transition from the typ i cal peri sphinctid, ancestral Jurassic Dorsoplanitinae to the Craspeditinae was accompanied by an extensive modification of the external septal suture to a characteristic and p ersistent craspeditid plan. The external and lateral lobes shortened, thickened at their bases, and became less incised. The retracted umbilical or suspensive lobe was replaced as a major element by a series of 4 to 7 auxiliary lobes arranged in order of steadily decreasing size away from the first lateral lobe towards the umbilical seam, sometimes so simplified as to be almost vestigial. The baseline is straight or only gently curved, and its sweep towards the umbilicus can be slightly adapical (retracted) or even adoral, sometimes called reversed. In the Polyptychitinae there was a partial return to the more usual perisphinctid plan. The family carries on consistently the style of di morphism established earlier. The micro conchs are isocostate with no adult peri stomal modifications of any kind, as in Dorsoplanites, making them often incon spicuous. The macroconchs are usually
18
Cephalopoda-Cretaceous Ammonoidea
variocostate, many becoming smooth, but there are isocostate exceptions (early Polypty chites) . The dimorphic size ratio lies between 2 : 1 and 4 : 1 . Relative abundances can be highly polarized, some known assemblages consisting alm ost completely of either macro- or microconchs. Ribbing is variable, but broad, short, subdued primary ribs or bullae are prominent up to the Valanginian. Most notable, however, is the frequent oc currence of polyschizotomous secondary rib bing in all combinations from dichotomous and bidichotomous to truly virgatotome, repeating the whole range found in what are thus seen to have been the ultimate ances tors, the Ataxioceratidae of the middle Up per Jurassic. This and the form of dimor phism distinguish Polyptychitidae from Olcostephanidae, with which they have at times been confused. Upper jurassic (Upper Volgian)-Lower Cretaceous (Upper Hauteri vian). Subfamily CRASPEDITINAE Spath, 1924 [ nom. transl SPATH, 1 9 3 1 b, p. 547, exCraspeditidae SPATH, 1 924c, p. [=Garniericeratinae SPATH, 1 952, p. 91
17}
Inner whorls compressed and involute; ribbing light and dense, soon fading; macro conchs tending to become inflated with re sidual, widely spaced, bullate primary ribs. Adults small to medium sized. Sutures highly simplified. Includes two separate series of faunas, one in the Spilsby Province, the other in the Russian Platform, Siberian Arctic, and Canadian Arctic p rovinces. GERASIMOV, 1 9 6 9 ; ] ELETZKY, 1 9 66; CASEY, 1 9 7 3 ; S HU L' GINA, 1 9 69a. Upper jurassic (Upp er Volgian)-Lower Cretaceous (Lower Rya zanian). Subcraspedites SPATH, 1 9 2 3c, p. 308 [*Ammonites plicomphalus J . de C. SOWERBY, 1 823, p. 145, non ]. SowERBY, 1 822, p. 82; SD SPATH, 1924c, p. 17; = S. sowerbyi SPATH, 1952, p. 1 8] . Inner whorls finely ribbed, wirh secondary ribs in dense sheaves pro jeered on venrer; outer whorls developing coarse, blum, shorr primary ribs. Sharpness of secondary ribbing external, nor impressed on inrernal molds. I n termediate between Laugeites and Craspedites. C ASEY, 1 973. Upper Jurassic (Upper Volgian)-Lower Cretaceous (Lower Rya:uznian): England. S. (Swinnertonia) S H U L'GINA, ! 972b, p. 1 3 8 [* Subcraspedites cristatus SwiNNERTON, 1935, p.
33; OD] . The earliest forms of Subcraspedites; evolure, densely ribbed like ancestral Laugeites (Up p er Jurassic); ribbing suddenly and strongly mo � fied in small adult macroconchs, giving widely spaced, sharp, primary ribs and dense sheaves of secondary ribs persisting to rhe end. CASEY, 1973; SWINNERTON, 1935. Upper Jurassic (Upper Volgian, primirivus Zone): England. F I G . 1 1 ,3a S. (Swinnertonia) subundulatus; roporype, macroconch, X 1 (Casey, 1 973; re prinred by permission of John Wiley & Sons, Lrd., © 1973). FIG. 1 1 ,3b,c. *S. (Swinnerto nia) cristatus; holorype, X 1 (Swinnerron, 1 9 35). S. (Subcraspedites) [= Volgidiscus CASEY, 1973, p. 239 (rype, Subcraspedites lamplughi SPATH, 1 936, p. 8 1 ; OD)]. Outer whorls moderately involute, primary ribs subdued, secondary ribs fine and dense, persisting to rhe end; adult body chamber beginning ro resemble Craspedites. UpperJurassic (Upper Volgian, preplicomphalus-lamplughi Zones): England. FIG. 1 1 , la-d. * S. (S.) sowerbyi SPATH; a, b, holorype, macroconch, X 1 ; c, X I , d, X 1 . 5 , roporypes (Casey, 1 9 7 3 ; re prinred by permission of John Wiley & Sons, Lrd., © 1973) . S. (Runctonia) CASEY, 1973, p. 240 [* R. runctoni; OD] . Involute, compressed microconchs wirh semifasrigiate venrer; secondary ribs chevronlike, fading on venrer; suture simplified on ascending baseline. Macroconchs apparently smoorh ex cept for ornamenr on umbilical and venrrolateral shoulders. Morphologically transitio n al ro Hectoroceras. Lower Cretaceous (Lower Rya:uznian, runcroni Zone): England.--FIG. 1 1 ,2a-d. * S. (R.) runctoni CASEY, holorype; a, b, shell, X 1 ; c, d, section, X 1 ; e, surure, X l . 5 (Casey, 1 973; re prinred by permission of John Wiley & Sons, Lrd., © 1 973). Craspedites PAVLOW, 1 892, p. 474 [non "Craspedites (pro - donites HAECKEL, ! 865) ALLMAN, 1 872, p. 1 74," cited by NEAVE, 1 939, p. 860, bur no trace of Craspedites in ALLMAN (hydro ids), hence presumably an incorrect subsequenr spelling of Craspedonites without s rarus] [*Ammonites okensis ORBI GNY, 1 845b, p . 436; SD R. DouviLLE, 1 9 1 1 , fiche no. 2 1 3] . Inner whorls compressed, involute, finely and densely ribbed; macroconchs becoming inflated, umbilicus narrow and craterlike, ribbing becoming rounded and subdued or fading altogether. Some species wirh constrictions. Upper Jurassic (Upper Volgian): England, Russian Platform, Arctic. C. (Craspedites) [=Mosquites SAZONOVA, 1 9 7 1 , p. 30 (type, Craspedites mosquensis GERASIMOV, 1960, p. 1 7 1 ; OD); Trautscholdiceras SAZONOVA, 1 977, p. 90 ( type, Amm onites kaschp u rensis T RAUTSCHOLD, 1 8 66, p. 1 6 ; O D ) ; Vitaliites SHU L'GINA, 1 9 8 5 , p. ! 1 6 (type, Ammonites subditus TRAUTSCHOLD, 1 877, p. 392; OD)]. Highly variable, usually involure and inflated, wirh smooth outer whorls; simple sutures rypi cal of subfamily. [Mosquites (for small cadicones retaining subdued secondary ribbing ro rhe end bur losing primary ribs) and Trautscholdiceras (for small cadicones with constrictions and with -
.
--
--
Ammonitina-Perisphinctaceae
19
Subcraspedites
I I
I I I
:
I I I I I '
3b
\' 2c
'
......
Runctonia
.
�-""· y
Swinnertonia
..
3c
FIG.
I I . Polyprychiridae (p. 18)
smooth body chambers except for well-spaced circumumbilical bullae) seem unnecessary.] GERASIMOV, 1969; SHUL'GINA, 1969a. Upper ju raSJic (Upper Volgian}: England, Russian Plat form (Moscow and Volga basins, Pechora), northern Urals, northern Siberia, Novaya Zemlya, Spitsbergen. FIG. 1 2,2a-c. * C. (C.) okemis (ORBIGNY). Russia; lectotype, macro conch, X 1 (R. Douville, 1 9 1 1 ) . C . (Taimyroceras) BOOYLEVSKII, 1956, p . 8 2 [• T taimyrmu; OD] Has most of rhe characters of C. (Craspediw), bur venter flatter with smooth band. [A large collection from a single concre tion gives a good idea of intraspecific variabiliry --
.
(jELETZKY, 1966).] ERSHOVA, 1969. Upperjuras sic (Upper Volga i n, taimyrensis Zone}: northern
Siberia, Sp itsbergen, Arctic Canada (Ellesmere lsland). FIG. 1 2 , Ja-c. * C. (T) taimyremis BODYLEVSKII, Kheta basin, northern Siberia; roporype?, macroconch, X I (Shul'gina, 1969a). C. (Kachpurites) SrATH, 1923c, p. 307 ['Ammonites folgms TtIAS in MUNIER-CHALMAS & DE LAPPARENT, 1894, p. 464 ('Ammonites verrucosw O RBIGN Y, 1841, p. 191; 00]. Dwarf, involute; in ner whorls inflated, smooth or with weak ribs; ourer whorls trapezoidal in section, with variable, sharp lateral and ventrolateral tubercles, opposite or alter nate on venter, appearing suddenly by thickening of ribs. Aperture of microconchs with long lappets. Suture with wide, short elements. THIEULOY, 1965. Lower Cretaceous (Lowtr Valanginian-Upper Valan ginian}: France, Spain, Balearic Islands, Germany, Mexico .--F IG . 34,4a,b. • S. vtrrucosum (O R BIGNY), France; XI (Failor & Termier, 1923). Parastieria S P ATH, 1923b, p. 144 [• Acantoceras? peltoceroides PAVLOW, 1892, p. 510; 00]. Dwarf; inner whorls as in 0. (0/costephanus), with 2 or 3 secondary ribs arising from slight umbilical bullae; outer whorl high and compressed, with straight or slightly sinuous, distant, high, Aat-ropped ribs and no tubercles. Aperture of microconchs with long, spatulate lappets. Lower Cretaceous (Lower Hauttri vian}: England.--FIG. 3 4, 3a, b. • P peltoceroides (PAVLOw); XI (Pavlow, 1892). Capeloites LISSON , 1937, p. 3 [*C. larozai; OD; =A m monites perelegans M ATHERON , 1878, pl. C-21, fig. 3]. Dwarf; outer whorl compressed with narrow, rounded venter, in early whorls umbilical bullae pinched, with coarse and fine ribs branching irregu larly; in later whorls umbilical bullae weakening and all ribs fine; aperture of microconch collared with broad, spatulate lappets. Lower Cretaceous (?Lower Hauterivian}: France, Peru.--FIG. 34 ,2a,b. *C. perelegam (MAT HERON) , Peru; X I (Thieuloy, 1969). Ceratoruberculus IMLAY, 1938, p. 568 [*C. casitensis; 00]. Dwarf; inner whorls as in Olcostephanus; outer whorls high, compressed, with prominent umbilical and ventrolateral bullae joined by bundled, looped, or zigzagged ribs; venter more or less grooved. Lower Cretaceous (Lower Hauterivian): Mexico.--FIG. 34,5 a, b. ·c. casitensis; XI (Imlay, 1938).--FIG. 34,5c,d. C. /inguituberculattiS IMLAY; XI (Imlay, 1938). Valanginites KI LI A N , 191 Oc, p. 196 ['Ammonites r�ucleus F. A. ROEMER, 1841, p. 87 (non PHILLIPS, 1829, p. 174; see ICZN Opinion 1610, 1990); SO ROMAN, 1938. p. 386] [=RotunditesSTOLLEY, 1937,
47
Ammonitina-Perisphinctaceae '
Olcostephanus
FIG. 32. Olcostephanidae (p. 46) p. 450, nom. nud.; Dobrodg�iaras NIKOLOV, 1963, p. 94. nom. nov. pro Dobrodg�ius NIKOL.OV, 1962, p. 69, r1on KrrrL, 1908, p. 522 (type, Dobrodgrites vmtrotflberculatus NIKOI.OV, 1962. p. 70; OD; =Holcosupbanus wi/fridi K;dlAKASCH, 1902, p. 14)]. Small, very involute sphaerocones with elliptical umbilicus and oblique, constricted, and collared ap erture; ornament ranging from very weak, straight, bundled or bidichoromous ribs with or wirhour slighr umbilical bullae to coarse, blunt, concave pri mary ribs branching at midside, with or without a large rubercle, into 3 to 6 strong secondary ribs; variably wirh (Dobrodgdctras} or without large siphonal tubercles on some ribs on later pan of body chamber. THIEULOY & GA7�Y, 1967; RICCARDI & WESTERMANN, 1970. Low�r Crttaetous (Lower
Valanginian-Upper Valanginian): France, Germany, Swirzcrland, Bulgaria, Ukraine (Crimea), Peru, Ar genrina.--FIG. 34, Ia, b. • V. nucl�m (ROEMER), Upper Valanginian, Germany; X I (Koenen, 1902). --FIG. 34,Jc,d. V. wi/fridi (I,
{ ' '
1b
!-..
Hengestites 1c
FIG. 97. Placenticeratidae (p. 126-130)
(type, P. murphyi; OD); Karamaites M. I. SoKOLOV in CASEY, 1965, p. 46 1 (M. I. SoKmov, 1961, p. 152, nom. nud.) (type, Placenticeras? grossouvrei
SEMENOV, 1899, p. 97; OD); Karamaictras M. I. SoKOLOV, 1967, p. 138 (type, K kolbajmse; OD); Trakmenites IL' IN , 1975, p. 154 (type, Placenticeras gouerdakemis LUI'I'OV, 1963, p. 144; 00); Kopetdagites lL'IN, 1975, p. 159 (type, K. kopetdagemis; OD); MediasicerasiL'iN, 1975, p. 159 (type, M. beliakovae; OD); Beschtubeites I!.'iN, 1975,
p. 162 (type, B. beschtubensis; OD); Asiatostantono ceras lL'IN, 1975, p. 172 (type, Stantonoceras tagammse; 00); Baghiceras CHIPLONKAR & G HARE, 1976, p. 3 (type, B. ambai; 00); Malwiceras CH!I'LONKAH & GHARE, 1976, p. 4 (type, M. variabilis; OD); Placentoscaphites CHII'LONKAR & G HAHE, 1977, p. 68 (type, P. dangerfeldi; 00); Sancarlosia CHIPI.ONKAR & GHARE, 1978, p. 79 (type, Placenticeras sancarlosense HYArr, 1903, p. 200; OD)]. Populations show great variability,
128
Cephalopoda-Cretaceous Ammonoidea
ranging from compressed, involute forms with con vergent, slightly convex sides, narrow, tabulate ven ter, no to weak, falcoid ribs, long umbilical bullae, and inner and more numerous outer ventrolateral tubercles to forms with similar inner whorls, but outer becoming more evolure, rectangular ro square in section, with more prominent umbilical and ven trolateral tubercles. the inner of which, originally midlateral, may move outwards to form the ventro lateral margin while the outer may disappear. In any population macroconchs comprise the larger, more compressed, and more feebly ornamented forms. The relative size of sutural elements varies slightly with time; early species tend to have the 4th lateral lobe smaller than the 5 th. Placenriceras was formerly distinguished from Karamaius by progressive in crease in number of sutural elements, by develop ment of strong dimorphism with strongly orna mented microconchs, and by progressive outward migration of umbilical tubercles to midlateral and of midlaceral to inner ventrolateral position. How ever, strongly ornamented microconchs are now known from Cenomanian and there seem ro be no valid grounds for dividing rhe slowly evolving se ries. Lower Cretnceous (Upper Albian)-Upper Creta-
ceous (Lower Campanian): France, Spain, Germany, Poland, eastern Africa. Madagascar, Israel, central Asia, southern India, British Columbia, USA, Mexico, Colombia.--FIG. 98a,b. P. grossouvrti SEMENO V , Cenomanian, Mangyshlak; X0.75 (Semenov, 1899).--FIG. 98c. P. mediasiaticum (Lurrov), Cenomanian, central Asia; X2 (Marci nowski, 1980).-- FIG. 99a.b. •p placenta (DE KAY), Campanian, New Jersey; X1 (Hyatt, 1903). --FIG. 99c-e. P. fritschi GRossouVRE, Coniacian, France; XI (Grossouvre, 1894).-- FIG. 99�h. P. guadaloupae (ROEMER), Campanian, Texas; X0.5 (F. A. Roemer, 1852).-- FIG. 99i,j. P. bidorsatum (ROEMER), Lower Campanian, Germany; X0.75 (MUller & Wollemann, 1906). Metaplacenticeras SPATH, 1926a, p. 79 [' Placenticeras pacificumSMITH, 1900, p. 187; OD] [=Paraplacen ticeras MATSUMOTO, 1953, p. 149 (type, Pfacmti cerassubtifistriatum)IMBO, 1894, p. 25(171); OD]. Compressed, flat-sided, with weak ro strong, falcoid m falcate ribs; veneer cricarinace, at lease in middle growth. Suture line follows char of ribs. SMITH, 1900; MATSUMOTo, 1953. Upper Cretaceous (?Santo nian, Campanian): Japan, California. --FIG. 97 ,I a-c. •M. pacificum () . P. SMITH), California;
Placenticeras
FIG. 98. Placenciceraridae (p. 126-128)
Ammonitina-Hoplitaceae
129
Placenticeras
fiG. 99. Placenticeratidae (p. 126-128) a,b, X0.75; c, X1 (Reeside, 1 926).--FIG. 97,Jd. M. JubtiliJtriatt<m QJMBO), Japan ; ?microconch, X 1
(Matsumoto, 1953). Hoplitoplacenticeras PAULCKE, 1 907. p. 183, ICZN Opinion 554 , 1 959, Generic Name No. 1348 (*HopliteJ plasticuJ PAULCKE, 1 907. p. 186; ICZN Specific Name No. 1629] (=Dechmoaras KAYSER, 1924, p. 175, nom. nud.]. Rather evolute; whorl
section compressed and parallel-sided to trapezoi dal; venter flat; with prominenr ribs varying from coarse and rounded to dense and fine; ribs nearly straight, with inner and outer venrrolatcral tu bercles, of which outer may be large and clavate; ribs crossing venter transversely and may have trace of siphonal tubercle. Uppa CruaceouJ (Uppa Campanian-Uppa Maastrichtian): France, The
1 30
Cephalopoda-Cretaceous Ammonoidea
Netherlands, Germany, western Africa, Egypt, South Africa (Natal), Madagascar, central Asia, British Columbia, Wyoming, Texas, Argentina (Pa tagonia).--FIG. 97,3a,b. • H plasticus PAULCKE, Campanian, Patagonia; X0.75 (Paulcke, 1907).
?Family ENGONOCERATIDAE Hyatt, 1900 (Engonocwridoe HYATT. 1900. p. 585] (=Knemiceraridae HYAlT, 1903,
p. 144; Neolobilinae LUPI'OV & MIKIIAILOV in LUJ'I'OV &. 0RUSHCHITS.
1958. p.
125]
Typically compressed, more or less flat sided, and involute, with venter Rat at least in some stage; single or branching, irregular ribs may occur with umbilical, lateral, or ventrolateral tubercles. Suture with numer ous auxiliary and adventive elements; saddles tending to simplify, being characteristically entire. More strongly ornamented forms giv ing rise to a succession of very compressed, smooth forms with narrow, bicarinate ven ters. Smooth offshoots are difficult to distin guish morphologically. Some genera closely resemble Pulchelliidae but are probably not closely related, and their origin remains un certain. HYATT, 1903; SPATH, 1923-1943; CASEY, 1960-1980. Lower Cretaceous (?Upper Aptian, Lower Albian)- Upper Cretaceous (Up per Cenomanian). Knemiceras BOHM, 1898. p. 200 [*Ammonitts syriacus BucH, 1 8 50, p. 20; OD] [=Glottouras HYATT, 1875, p. 372 (type, Buchicerm attmrEN, 1955c, p. 380 (rype, 0. � mtbu ndu m; 0D); ? utragonoceras HoEPEN, I 95 5c, p. 382 (rype, T. robustum; 00)]. Small, evolure, with square or rectangular whorl section; keel strong at firsr, weakening, and disappearing before end of last whorl; ribs single or branching, rypically sinuous, thickened on shoulders, and rending ro overhang backward. Lo wu Cretaceous (Uppu Albian): South Africa (Zululand).--FIG. 1 0 4,5. •£. unuis; X I (Hoepen, 1955c). Neokenrroceras SPATH, 1921a, p. 306, ICZN Opinion 1 2 54, 1 9 8 3 , Generic Name No. 2196 (*N. curvicornu SPATH, 1922a, p. 139 (1921a, p. 306, nom. nud. ); ICZN Specific Name No. 2863]. Small; outer whorls as in Hysttroceras; inner whorls with very strong umbilical tubercles; weak or no lateral tubercles; large, clavate, normally irregular, ventro lateral tubercles; and ribs weak or absent. Lower Crttaaous (Upptr Albian): western Africa, ?Bra ziL--FIG. 1 04,2a,b. N. curvicornu, Angola; in ner whorls, X I (Sparh, 1922a). *
Subfamily MOJSISOVICZIINAE Hyatt, 1903 [nom. trans/. WRICIIT. 19S2. p. 221, aMojsisoviaiidac HYATT, 1903, p. 241 (=Dlpoloccr:utdac SPAT! I, 192Ja, p. 2n; �chenoccr.uid;zc HOEPEN, 1941, p. 61; Dr<panoccr.uidac HOEPEN, 1941, p. 891
Derivatives of Brancoceratinae, in which the keel tends to become stabilized as an adult feature and the ribs begin to differen tiate. In some forms the keel appears early in ontogeny and is then lost; in others it only appears lace. Apparently, there are two main stocks (both derived from Mojsisoviczia or one from che other): one compressed and high-whorled, which left no descendants, and the other more evolute, with round or square whorl section, which by acquisition of tubercles led to Mortoniceratinae. The boundary between subfamilies is arbitrarily taken at the point when large umbilical and
137
ventrolateral tubercles are stabilized in the adult. Lower Cretaceous {Lower Albian-Upper Albian). Mojsisoviczia STEINMANN, 1 8 8 1 , p. 142 [*M. dunftl di; 00; =Ammonius vmtanill�mis GABB, 1 869, p. 273) [=Dipoloaroid�s BREISTROfFER, 1 947b, p. 90(74) (rype, Ammoniw d�laru�i ORBIGNY, 1841, p. 296; 00)]. Macroconchs up ro I 00 mm in diameter; evolure and smooth in young, with round whorl section; later with prominent, sharp keel and strong, straight or slightly curved, single ribs, which m�y expand in breadth on shoulders or form large m1dlareral and ventrolateral tubercles. Microconchs 25 to 35 mm in diameter; smooth at first, rhen with feeble ribs and slight to moderate keel; finally smooth and without keel bur generally with a few constrictions. The lecrorype (GEBHARD, 1983) of M. duufildi is a microconch here held ro be of the as sociated M. vmtanillmsis (GABS). KENNEDY & Coo PER, 1977; GEBHARD, 1 983. Lower Cretaaous (Middl� Albian}: Europe, South Africa (Zululand), Pakistan, Texas, Mexico, Colombia, Peru, Venezu ela, Greenland.--FIG. 105.l,a-e. • M. ventanil lmsis (GABS), Peru; a,b, lecrorype of M. duerftldi, XI (Gebhard, 1983); c-e, XI (Douglas, 1 9 2 1 ) . --FIG. l 05.1jg. M. d�laru ei (ORUIGNY), France; XI (Cooper, 1982). Falloriceras PARONA & BONARE LLI, 1897, p. 89 (•Am moniteJ proteus ORB IGN Y , 1842a, p. 624; 00]. Closely resembles the smooth microconchs of Mojsisoviczia, but early whorls have weak ribs and keel declines at maturity. More probably a progeneric dwarf ofsf hoot of Mojsisoviczia (KEN NEDY & COOI'ER, 1977), rather than a microconch of any species of Mojsisoviczia (GEBHARD, 1983). Lowu Cretac�ous (lower Middle Albian): England, France, South Africa (Zu!uland), Pcru.--FIG. . 105,2a-c. *F prouus (O RBI GN Y), France; a,b, X I ; c, X6 (Kennedy & Cooper, 1977). Oxytropidoceras ST!ELER, 1920, p. 346 [*Ammonites roissyanus ORBIGNY, 1841, p. 302; OD] [=Psrodo phacouras SPATH, 192la, p. 281, obj.]. Moderately ro very compressed and high-whorled, with high keel; ribs single ro branching, narrow and high, rounded or flat, with or without tubercles; keel may appear well before ribs; ribs may be effaced later. Suture generally with oblique outer slope on first lateral saddle. Lowu Cretaceous (Lower Albian-Up per Albian): Europe, Morocco, Angola, South Africa (Zululand), Madagascar, Pakistan, California, Texas, Peru, Colombia, Venezuela, Brazil, Mexico. 0. (Oxyrropidoceras) [=Manuaniaras SPATH, 1925c, p. 182 (type, Pseudophacoceras manua n�meS!'ATH, 192la, p. 281; 00)]. Moderately to very high-whorled and compressed; rather involute; ribs coarse to fine, high and narrow ro flat, mosrly branching at various levels, and without tubercles. Lowu Cr�taaous {Lower Albian-Middle Albian): Europe. Angola, South Africa (Zululand), Madagascar, Pakistan, Texas, Peru, Colombia, BraziL--FIG. I 06,2a-c. • 0.
138
Cephalopoda-Cretaceous Ammonoidea (0.) roissyanu m (ORBIGNY), France; a.b. X0.75; c. enlarged (Orbigny, 1 840-1 842).--FIG. 106,2d.�. 0. (0.) manttaneme (SPATH), Zulu land; XI (Spath, 192la).
0. (Mirapelia) CooPER, 1982, p. 29 1 ['Ammonites mirap�lianttm ORBIGNY, 1 8 50a, p. 1 24; 00]. Whorl section varying from only a little higher than wide to moderately compressed; ribs single, coarse to rather fine, and markedly Aattcncd and broadened on shoulders. Lower Cntauoru (Middle Albian}: England, France, Angola, Madagascar, California, Texas, Colombia, Peru, BraziL--FIG. 1 06 , Ia, b. * 0. (M.) mirapeli anttm (ORL\IGNY), France; X0.5 (Cooper, 1982). 0. (Venezoliceras) SPATH, 1 925c, p. 1 8 2 [* 0. 11m�zolanum STIELER, 1 920, p. 394; 00] [= Lophocaas HOEPEN, 1 93 1 , p. 40, non HYATT, 1 893. p. 466 (type, L. umsinmse; 00); Tarfizy ites COLLIGNON, 1967, p. 1 9 ( 1 963, p. 142, nom. nud.) (type. 0. (Tarfayius) bi tttbucu latu m; 00)]. Ribs straight or slightly sinuous, typically coarse, high, steep in front, shallow behind, but may be Aat at some growth stage; high umbilical to midlateral bullae on some ribs and bullate to slightly clavate ventrolateral tubercles on all.
Lower Cr�tauous (Midd/� Albia11-Upper Albian}: Morocco. South Africa (Zululand), Madagascar,
Texas, Mexico, Venezuela, Peru. --FIG. 107, 2a,b. •a. (V.) vmezolanum, Venezuela, nco type; X I (Renz, 1982). 0. (Laraiceras) RENZ, 1968, p. 650 [• 0. (L.) laramu; 00]. With alternate, long and short, coarse ribs; long ribs with umbilical, midlateral, and ventrolateral tubercles. Lower Cr�taceotts (Upper Albian}: Venezucla.--FIG. 107.3a.b. •a. (L.) laracnu; X0.5 (Renz, 1 982 ) . 0 . (Benavidesites) COOI'ER, 1 9 8 2 , p. 2 9 5 [• Vmezoliuras harrisoni BENAVIDES-CACERES, 1956. p. 460; 00]. Ribs fine and bifurcating at first, then coarsening, with midlateral tubercles on outer whorls. Lower Cretaceous (Middle A lb i an) : Texas, Venezuda, Peru. -- FIG. I 06,3 a. • 0. (8.) harrisoni (BENAVIDES-CICERES), Peru; X0.5 (Benavides-Caceres, 1956).--FIG. I 06,3b. 0. (B.) ac u toc arinatu m (SHUMARD), Texas; XO.S (K. Young, 1966). 0. (Adkinsitcs) SPATH, 1 93 l a, p. 350 ['Ammonites belknapi MARCOU, 1 8 58, p. 34; 00] [=Androiavites COLLIGNON, 1936, p. 186 (type. 0. (A.) bnairiei; 00)]. Ribs coarse, variable, straight or curved; inner whorls at least with strong umbilical and weak ventrolateral tu bercles. Lower Cretauous (Middle Albian-Upper Albian}: Europe, Madagascar, Texas.--FIG.
t �� . �y . �c r
Fallotice ras
Mojsisoviczia
1e
.
FIG. 105.
Brancoceratidae (p. 137)
Ammonitina-Acanthocerataceae
139
Oxytropidoceras Mirapelia 1a
FIG. 106. Brancoceratidae (p. 137-138)
107,1. 0. (A.) bravoeme (BOSE), Texas; X I (K. Young, 1966). Dipoloceras HYAIT, 1900, p. 589 ['Ammonites cristatus BRONGNIART in CUYlER & BRONGNIART, 1822 , pl. 0, fig. 9; OD]. Rather evolme; rypically inflated or
depressed; keel prominent, commonly below level of ventrolateral ends of ribs; ribs dense co distant, rounded co sharp, rypically a mixcure of single and branched, the latter flared at point of branching; umbilical and ventrolateral tubercles present in
140
Cephalopoda-Cretaceous Ammonoidea some. Sutures with plump, broad, finely indented saddles. Lower Cr�taceous (Middle Albian- Uppa Albian}: Europe, South Africa (Zululand), Mada gascar, Japan, Texas, ?Cura�ao. D. (Dipoloceras) [= Cechmoceras HOEPEN, 194 1 . p. 6 1 (type, C. revtrsum; OD)]. Whorl section round or depressed; umbilical and ventrolateral tubercles absent or insignificant. Occurrence and distribution as for genus. FIG. I 08,1a-c. • D. (D.) cristatum (B RO N G N I A RT in CUYlER & BRONGNIART), Middle Albian, France; a.b. X I ; c. X2 (Orbigny, 1840-1842). D. (Rhytidoceras) HOEPEN, 1 9 3 1 , p. 42 [*R. &gam; OD] [=Ricnouras HOEPEN, 1941, p. 59 (type, R. pandai; OD); Diplasioceras HOEPEN, 1946a, p. 203 (type, D. fa/lax; O D) ; Eusputro· aras HOEPEN. 1 946a, p. 202 (type, E. strigile; OD); 'Mortoniceratoides CooPER, 1982, p. 296 (type, Mortoniaras rigidum S PATH, 1933b, p. 4 13; OD)]. Inner whorls rending ro be Aar sided, wirh disrincr umbilical and weak venrro lareral tubercles and with ribs a mixture of branched and flexuous and single and straight; strong, spiral notching may occur (Euspectro· aras). Occurrence and distribution as for ge nus.--FIG. 108 2a, b . • D. (R.) elegans (HOEPEN), Upper Albian, Zululand; X0.5 (Hoepen, 1941). ?Menuthengonoceras COLLIGNON, 1963, p. 130 [*M. komihevitramu; OD]. Based on single fragment of discoidal, high-whorled form with shouldered ven ter and high keel. COLLIGNON (1963) compared rhe sucure of Mmuthengonoaras with Hypengonoaras, bur rhe fragment seems as likely to belong to Mojsisovicziinae as to Engonoceraridae. Lowa Cre· taceous (Middle Albian}: Madagascar. --
,
Subfamily MORTONICERAT INAE H. Douville, 1 9 1 2 [nam. trans! S!>ATII. 1934, p . 465. tx Mononiccratidac SPATH, 1925c. p. 182, uom. rorr((J. pro MononiccracinCs H. OoUVILL£, 1912, p. 295] [=lnflaticcrarid.:a� SPATII, 1925c. p. 181: Pervinquicriid2e SrAru, 19263, p. 79: Arestou:r.uidae HOEPtN, 1942, p. 1 17; Cainocc.r:uidac HoEPEN, 1942, p.
Venezoliceras FIG. I 07. Brancoceraridae (p. 138-139)
2a
1271
Moderately involute to very evolute; whorl section more or less rpunded, square, or compressed, with low to high keel; ribs branching or long and short, at least in early whorls, bur may be single and equal on body chamber or earlier; ribs low and rounded, or flat, or high and rounded, but not high and sharp; at least umbilical tubercles occurring, normally also inner ventrolateral, and fre quently lateral and outer ventrolateral; some species with up to 5 tubercles on a rib; spi ral striation or notching of ribs common; aperture simple (in probable macroconchs) or with rostrum directed forward, upward, or backward (in probable m i croconchs).
Ammonitina-Acanthocerataceae Suture generally with squarish, symmetrical, deeply and sharply indented saddles. HOEPEN, 1 94 1 - 1 9 5 1 ; SPATH, 1 9 23-1943. Lower Cretaceous (Middle Afbian)-Upper Cretaceous (Lower Cenomanian). As the synonymies of subfamily and gen era show, both the nomenclature and scale of classification of the group have been in doubt. A large number of species with many of the same basic characters vary in detail and in combination of whorl section, num ber and shape of tubercles at different growth stages, and in strength and direction of ribs. An attempt is made here to classify them in accord with the scale accepted i n allied groups. The subfamily is derived from Mojsisovicziinae along one or several closely allied lines and remains very uniform. Mortoniceras MEEK, 1 876, p. 448 ['Ammonites tmpertinus MoRTON, 1834, p. 40; 00 (despite many authors, a valid species)) [for synonyms see subgenera]. More or less evolute, with square, rect· angular, or trapezoidal costal whorl section; ribs normally strong bur may weaken or strengthen on body chamber; with prominent umbilical and nor mally with inner ventrolateral tubercles at least and, ar some growth stage, I or 2 lateral and an outer ventrolateral tubercle also may occur; keel high or low. Lower Cretaceous (Middle Albian-Upper Albian}: Europe, Africa, India, North America, South America, Madagascar. M. (Morroniceras) [=Pervinquieria BOHM, 1910, p. 1 52 (type, Ammonites inflaws ) . SoWERllY, 1 817c, p. 170; M); lnflaticeras STIELER, 1 920, p. 346 (type, Ammonites inflatus J. SowERBY, 1 817c, p. 170; 00); Subschloenbachia Si'ATH, 1 9 2 1 a, p. 284 (type, Ammonites rostratus J. SowERBY, 1 817b, p. 163; 00); Leoniw SP!ifH, 1932, p. 388 (type, Ammonites leonensis CON· RAD, 1857. p. 160; 00); Ophryoceras HOEI'EN, 1942, p. 9 1 (type, 0. jugosum; 00); ?Ameleceras HOEI'EN. 1942, p. 1 1 5 (type, A. abjectum; 00); Rusoceras HOEPEN, 1946a, p. 238 (type, R. nothum; 00); Collignonia HOEPEN, 195lb, p. 295 (type, Pervinquieria (C.) undulatocoslata HOEI'EN, 1 95 l b, p. 295; 00); Styphloceras HOEPEN, 1951 b, p. 300 (type, Pervinquieria (5.) nodosocostata HoEPEN, 1 9 5 1 b, p. 300; 00); Omocrateceras HoEPEN, 1 9 5 l b. p. 3 1 3 (type, Pervinquieria (0.) planiventer HoEPEN, 1 9 5 1 b, p. 313; 00); Subpervinquieria MIRZOEV, 1969. p. 46 (type, Pervinquieria (S.) luppovi MIRZOEV, 1 969, p. 46; 00)). Ribs moderately fine to very coarse, normally branching at umbilical tu· bercles on early whorls, later single; with umbili cal and inner ventrolateral tubercles; mediolat· era! tubercles typically weak, present or not; outer ventrolateral tubercle commonly percep-
141
2b
Rhytidoceras FIG. I08. Brancoceratidae (p. 140)
142
Cephalopoda-Cretaceous Ammonoidea rible at some stage and in some species, e.g., M. (M.) rostratum (]. Sowmuv), and may be strong for a short stage, foreshadowing M. (Dumova n"tes); in a few species all tubercles weak and ribs dominant; rostrum (questionably of micro conchs) raised and straight ro recoiled. Occur rence and distribution as for genus except rhar it does nor reach rhe extreme rop of rhe Albian.
--FIG. 109a-d. ·M. (M.) vnpertinum (MOR· TON), Upper Albian, Texas; a,b, holorype, X0.25; c,d, X0.5 (new).--FIG. 109 e-g. M. (M.) inflatum {). SOWERBY); e, Upper Albian, England, X0.7; f. Upper Albian, England, X I (Spath, 1923-1943); g, Upper Albian, France, XI (Orbigny, 1840-1842).--FIG. 109h. M. (M.) rostratum () . SOWERBY), Upper Albian,
France; X0.5 (Scholz, 1979). (Boesires) K. YOUNG, 1 9 6 8 , p. 7 1 [• Pervinquitria romeri HAAS, 1942, p. 73; OD]. Very evolure, with umbilical and ventrolateral bullae bur no lateral tubercles; ribs strong and distant, equal or long and short, bur nor bifur cating. Barely distinct from some biruberculare
M.
M. (Mortoniams). Lower CrNaceous (Upper Albian): Angola, Texas, Mexico. M . (Deiradoceras) H O EI'E N , 1 9 3 1 , p . 52 [*lnjlaticeras prcrostratum SPATH, 1 9 2 l a , p. 284; OD] [=Drepanoceras HOEPEN, 193 1 , p. 46, non STEIN, 1 8 7 8 , p. 25 (type. D. undatum; OD); Mimeloceras H OEI'EN , 1944, p. 196, nom. nov. pro Mimoceras HOEPEN, 1941, p. 85, non HYATT, 1 88 4 , p . 309 (type, Mimoceras binodosum;
OD)]. Whorl section depressed ro slightly com pressed, subquadrare, with sharp keel; very strong umbilical and ventrolateral tubercles with cosral whorl section concave between; midlateral tubercles may be visible at some stage; early whorls rending ro have rounded venter as in Dipolouras; rostrum projected in concinuarion of coiling of shell. Lower Craauotts (Upper Albian): western Europe, Africa, Venezuela.- FIG . I I 0,3. •M. (D.) pmostratum (SPATH), South Africa (Zululand); X0.75 (Spath, 1 921a). M. (Durnovarites) SPATH, 1932, p . 380 [* Subschlombachia perinjlata SPATH, 1922a, p. 1 1 3; M; SPATH ( 1932, p. 380) designated the type as follows: "M. (D.) subquadratum, nov. Plate xxxvii, figs. 6a,b (Diagnosis below)"; how ever, the diagnosis did nor appear until 1933 (p. 429) and, after 1931, mere reference to a figure was insufficient to validate a species. Thus, M. (D.) subquadratum was in 1932 a nom. nud. SPATH ( 1933b, p. 429) quoted the type species of Du rnovarites as M. (D.) perinjlatum (S PATH ) without comment.] [=Reyreiceras COLLIG NON , 1979, p. 34 (type, R. reyrei; OD)]. Whorl sec tion square ro depressed-trapezoidal; from early stage at least 4 nearly equally spaced tubercles on each rib, the outer one as prominent as the oth ers and rending ro be clavate. Lower Cretauous (Upper Albian): Europe, Africa, Madagascar, Texas.--FIG. I I 0, 2a, b. 'M. (D.) perinjlatum (SPATH), France; ?microconch, X0.5 (Scholz,
1 9 7 9 ) . - FIG. L I 0 ,2c,d. M. (D.) subqua dratum, England; XL (Spath, 1 923-1943). M. (Angolaires) SPATH, 1932, p. 380 [* Pervin quieria gregoryi SPATH, 1922a, p. 1 27; OD] [=?Praeangolaites COLLIGNON, 1 979, p. 41 (type, P galvaoi; OD)]. Rather compressed; with single
ribs at all stages; at first with umbilical tubercles; later also with prominent inner and outer ven trolateral tubercles. Lower Cretaceous (Upper Albian): western Africa.--FIG. I I 0, Ia, b. •M. (A.) grtgoryi (SPATH), Angola; X0.5 (Spath. 1922a). M. (Drakeoceras) K. YOUNG, 1957, p. 19 [*D. drakei; OD]. Compressed to rather inflated; normally wirh branching ribs at some stage and with double ventrolateral tubercles as in M. (Angolaius). Lowtr Cmaaow (Upptr Albian):
Angola, Texas.--FIG. I I 0,5a,b. *M. (D.) drakei (YOUNG), Texas; X0.75 (K. Young, 1957). M. (Pagoceras) HOEPEN, 1 9 5 1 b, p. 324 [� P amplijicatum; OD] [=Poikiloceras H O EPE N , 1951 b, p. 329 (type, P Jirmum; 00)]. Moder ate-sized, with markedly fastigiate venter becom ing increasingly acute with age. Lower Cretaceous (Upper Albian): South Africa (Zululand).- FIG. I I0,4a,b. •M. (P) amp/ijicatt<m
(HOEI'EN);
(Hoepen, 1 9 5 1 b). Goodhallites SPATH, 1932, p. 3 8 1 [•Ammonites good halli J. SowERBY, 1820, p. 100; 00] [=Letlmaas HOEI'EN, 1 942, p. 138 (type, L. complanatum; 00); Cainoceras HOEPEN, 1 942, p. 149 (type, C. liberum; 00); Lethargecems HOEI'EN , 1942, p. 149 (type, L. incommodum; OD); Aidoaras HOEPEN, 1946b, p. 248 (type, A. jubatttm; O D) ]. More or less com pressed and high-whorled, with high keel·; inner whorls generally finely ribbed, with rather weak umbilical, venrrolareral, and rarely midlareral tu bercles or none; outer whorls normally with strong ornament, tubercles becoming more prominent with age, bur final part may be nearly smooth. Lowa Cretaceous (Uppa Albian): Europe, Africa, Madagascar, southern India, Australia (Queens land), Texas, Venezuela.--FIG. 1 1 1 , 3a, b. ·c. goodhalli (J. SowERIIY), England; a, X0.67; b, X0.5 (Spath, 1923-1 943). --FIG. 1 1 1 ,3c,d. G. fiber (HOEPEN), South Africa (Zululand); X I (Hoepen, X0.5
1942).
Arestoceras HOEI'EN, 1942, p. 1 1 8 ['A. collinum: OD] [=? utagmmoceras HOEPEN, 1942, p. 122 (type. T. tumu/omm H O EPF.N , 1942, p. 123; SD WRIGHT. herein)]. Early whorls as in some early, less rubercu lare species of M. (Mortoniceras), bur ornament rending to disappear, whorl height to increase, and venter to become fastigiate rather than keeled. Per· haps a subgenus of Goodhallites. Lower Cretaceous (Upper Albian}: South Africa (Zululand).--FIG. I l l ,2. •A. collinum; XO.75 (Hoepen, 1942). Neoharpoceras SPATH, 1 92 1 a, p. 282 [•Ammonim hugardianus ORBIGNY, 1 84 1 , p. 2 9 1 ; 00]. Com pressed; more involute than Prohystaoceras and wirh more rounded venter and lower keel; ribs dense and sinuous, without tubercles. Suture florid. Lower Cretaceous (Upper Albian): western Europe.--FIG.
Ammonitina-Acanthocerataceae
FIG. 109.
Brancoceraridae (p. 141-142)
143
144
Cephalopoda-Cretaceous Ammonoidea
Drakeoceras FIG. 1 10. Brancoceratidae (p. 142)
hugardianum (ORBIGNY). France; (Orbigny, 1840-1842). Prohysteroceras SI'ATH, 1921a, p. 286, 1CZN Opinion 1254, 1983, Generic Name No. 2195 ['?. worditi SPATH, 1922a, p. 143 (1921a, p. 286, nom. nud.); ICZN Specific Name No. 2862]. Rather evolute I I I ,la,b. • N.
X0.75
and moderately compressed; ribs fine, close, sinu ous, and branching; tubercles weak or absent; ros rrum prominent, directed upward. Lou11:r Cretactous (Upptr Albian): England, Angola. FIG. 1 1 1,4a c. •P. worditi, An gola; tt, X0.7; b, X I ; c, X2.5 (Spath, 1922a). --
Ammonitina-Acanthocerataceae
Neoharpoceras
FIG. I l l . Brancoceratidae (p. 142-144)
145
146
Cephalopoda-Cretaceous Ammonoidea
}1
6
Elobiceras
FIG. 1 12. Brancoceraridae (p. 146-147) Elobiceras SI'ATH, 1 92 1 a, p. 306 (*Schloenbachia �lobiensis SZAJNOCHA, 1885, p. 235; OD]. Rarher evolure and compressed, wirh numerous clavi or spiral norchcs on rhe ribs. Lowa Cretactotts (Upper Albian): Angola, sourhern India, Nigeria, Texas. E. (Craginires) K. YoUNG, 1957, p. 14 (* Schloen bachia leonmsis var. serrattscens CRAGIN, 1893, p. 2 4 1 ; O D ) . Wirh alrernare, long and shorr, flexuous ribs. Occurrence as for genus: Texas. --FIG. 1 1 2,5. • £. (C.) serrattscms (CRAG IN) ; XI (K. Young, 1957). E. (Eiobiceras). Ribs single, recriradiare. Occur rence as for genus: Angola, Nigeria. southern lndia. --FIG. 1 1 2, 6. *E. (E.) rlobiense (SZAJNOCHA), Angola; X0.5 (Szajnocha, 1885). Cantabrigites SI'ATH , 1933b, p. 436 ( 1 932, p. 380, nom. rmd.) [•Mortoniceras (C.) cantabrigense SPATH,
1932, p. 380; OD]. Small; generally with branch ing ribs ar firsr, rending ro become single, mosrly
nontuberculare. Suture much simplified. May be closer ro Hysteroaras (Brancocerarinae). Lower Cre taceous (Upper Albian}: England, France, Swirzer land.--FIG. 1 12,1 a-c. •c. cantabrigmse SPATH , England; a,b, X I ; c, X2 (Sparh, 1 923-1943). Algericeras SPATH, 1925c, p. 182 [•Ammonites boghar i COQUAND, 1880, p. 35; OD] [=Prionocycloides imss Si•ATH, 1925c, p. 1 8 2 (rype, Ammonites proratus CoQUAND, 1880, p. 32; OD)]. Small; compressed ro inAared, wirh rounded shoulders and keel; ribs fine or coarse, sharp and branching. Seems ro be derived from Cantabrigites. KENNEDY & WRIGHT, 1 9 8 1 . Uppa Cretaceous (Lower Cenomanian): norrhern Africa, Madagascar, Mexico. A. (Algericeras). Keel nodare ar firsr, rhen entire. Occurrence as for genus: norrhern Africa.- FIG. 1 1 2,3. *A. (A.) bogharimse (COQUAND), Tunisia: X2 (Pervinquiere, 1907). A. (Sakondryella) COLLIGNON, 1964, p. 2 1
Ammonitina-Acanthocerataceae [*Euhystrichouras (S.) madagascarieme CoLLIG NON, 1 9 6 4 , p. 2 1 ; 00; =Euhystrichouras remolineme BOSE, 1 928, p. 247]. Keel wich
strong crenulations throughout. Occurrence as for genus: Madagascar, Mexico. FIG. 1 1 2,2a, b. •A. (S.) mnolinmu (BOsE), Madagas car; XI (Collignon, 1964). Euhysuichoceras SPATH, 1923b, p. 143 [•AmmoniUs nicaiui COQUAND, 1 862, p. 323; 0 0 ] . Small; rather evolute; compressed co inflated, wich Rat or convex sides and strong, emire keel; irregular ribs springing in cwos and threes from sharp umbilical tubercles and curving forward on vencer; shorter ribs intercalated; vencrolareral tubercles may occur and if so !auriform ribbing may develop. KENNEDY & WRIGHT, 1 98 1 . Upptr Cretaceous (Lower Cmo manian}: England, northern Africa, Nigeria, Mada gascar, Mexico. FIG. I I 2,4a, b. ·E. nicaiui (Co QUAND), Tunisia; X I (Pervinquiere, 1907). --
--
Family LYELLICERAT IDAE Spath, 1921 [Lyellie memoir43:1-24, 5 fig., pl. 17-19. ---. 1 9 1 1 . AmmoniteJ okmsis d'Orbigny 1 8 4 5 . Palaeontologia Universalis (series 3 ) fascicle 3, fiche 213. ---. 1 9 1 2a. [Review of "Untersuchung einiger Ammonitiden aus dem unteren Gault Mangyshalak.s und des Kaukasus," by J. Sinzow, 1908]. Revue Cri tique de Paleozoologie 1 6:260. ---. 1 9 12b. Etudes sur les cardiocCtatides de Dives, Villers-sur-mer et quelques autres gisements. Memoires de Ia Societe Geologique de France, Paleontologie 19, memoir 45: 1 -77 , fig. 1-84, pl. 15(7- 1 1 ) . Doyle, J. C . 1 963. A new hereromorph ammonite from the Lower Cretaceous of Yorkshire. Palaeontology 6,575-578, £g. I , pl. 78. Drushchits, V. V. 1953a. K izucheniia ontogenet icheskogo razvitiia Biosaloceras subuquens (Karakasch) [Study of the onwgeneric development of Bid.Saloceros subsequens (Karakasch)]. Vestnik Moskovskogo Univmiteta, no. 6 : 1 57-1 6 1 . ---. 1953b. Dannye o b ontogeneze novogo roda ammoniwv Euphylioceras [Data on rhe ontogeny of rhe new ammonite genus Euphylloceras] . Vestnik Moskovskogo Universiteta, no. 9 : 1 4 1 -147. ---. 1956. Nizhnemelovye Ammonity Kryma i severnogo Kavkaza [Lower Cretaceous Ammonites, Crimea and Northern Caucasus]. Izdatel'stvo Moskovskogo Universiteta. Moscow. 149 p., 13 pl. ---. 1960. Ammonitii [Ammonites], p. 249-355, pl. 1-47, 1-22. In V. V. Drushchits & M. P. Kudriavtseva, eds., Atlas Nizhnemelovoi fauny severnogo Kavkaza i Kryma [Ada.� of the Lower Cre taceous faunas of the northern Caucasus and the Crimea] Vsesoiuznyi Nauchno-Issledovatel'skii Institut Prirodnykh Gasov. Moscow. Dubourdieu, Georges. 1953. Ammonites nouvelles des Monts du Melll!gue. Bulletin d u Service de Ia Carte Geologique de l'AJgerie (serie 1, Paleontologie) 1 6 : 1 76, £g. 1-20, pl. 1-4. Dujardin, M. J. L 1837. MCmoire sur les couches du sol en Touraine, et description des coquilles de Ia craie et des faluns. Memoires de Ia Societe Geologique de France (series 1 ) 2(1835):2 1 1-3 1 1 , pl. 1 5-20. Dumas, Emilien. 1876. Statiscique Geologique, Mineralogique, MCtallurgique ct Paleontologique du Department du Gard, part 2. Paris & N!mes. 735 p., 9 pl. Durand Delga, Michel. 1954. A propos de "Bochianitrs'" supentes Perv. : Remarques sur les ammonites droites du Cretace infCrieur. Compte Rendu Sommairc des Seances de Ia Societe Geologique de France 1954,134-137. I £g. Egoian, V. L. 1 9 6 5 . 0 nekotorykh ammonitakh klanseia zapadnogo K.avkaza [Some ammonites of the Clansayesian of the western Caucasus]. Trudy Krasnodarskogo Filiala Ysesoiuznogo Neftegazovogo Nauchno-Issledovatel'skogo lnstituta 16: 1 1 2-160, 14 pl. ---. 1 9 6 9 . Ammonity iz klanseiskikh sloev zapadnogo Kavkaza [Ammonites of the Clansayesian beds of the western Caucasus], p. 126-188, 26 pl. In V. L, Egoian, cd., Geologiia i neftega:zonosnosr'
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