Bulletin 64

New Mexico Museum of Natural History & Science

A Division of the DEPARTMENT OF CULTURAL AFFAIRS

CONODONTS FROM THE CARNIAN-NORIAN BOUNDARY (UPPER TRIASSIC) OF BLACK BEAR RIDGE, NORTHEASTERN BRITISH COLUMBIA, CANADA

by MICHAEL J. ORCHARD

Albuquerque, 2014 Bulletin 64

New Mexico Museum of Natural History & Science

A Division of the DEPARTMENT OF CULTURAL AFFAIRS

CONODONTS FROM THE CARNIAN-NORIAN BOUNDARY (UPPER TRIASSIC) OF BLACK BEAR RIDGE, NORTHEASTERN BRITISH COLUMBIA, CANADA

MICHAEL J. ORCHARD

New Mexico Museum of Natural History & Science Albuquerque, 2014 STATE OF NEW MEXICO Department of Cultural Affairs Veronica Gonzales, Secretary

NEW MEXICO MUSEUM OF NATURAL HISTORY AND SCIENCE Gary Romero, Interim Executive Director

BOARD OF TRUSTEES Susanna Martinez, Governor, State of New Mexico, ex officio Charles Walter, Executive Director, ex officio Gary Friedman, President Deborah Dixon Maya Elrick, Ph.D. Peter F. Gerity, Ph.D. Laurence Lattman, Ph.D. Morton Lieberman, Ph. D. Imogene Lindsay, Emerita Viola Martinez Marvin Moss John Montgomery, Ph.D. Jennifer Riordan Laura Smigielski-Garcia David Smoak Steve West

Cover illustration: Some evolving lineages of conodonts from the Carnian-Norian boundary beds: 1-3. Metapolygnathus ex gr. communisti> M. dylani> M. parvus; 4-6. Parapetella riteri beta> P. riteri alpha> P. willifordi; 7-8. Acuminatella sagittalis> A. acuminata.

Original Printing ISSN: 1524-4156

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NMMNH Bulletins online at: http://nmnaturalhistory.org/bulletins BULLETIN OF THE NEW MEXICO MUSEUM OF NATURAL HISTORY AND SCIENCE

EDITORS

Spencer G. Lucas New Mexico Museum of Natural History and Science, Albuquerque, NM, USA (NMMNHS) Robert Sullivan NMMNHS Lawrence H. Tanner Le Moyne College, Syracuse, NY, USA

PRODUCTION EDITOR

Asher J. Lichtig NMMNHS

ASSOCIATE EDITORS

Guillermo Alvarado Asociación Costarricense de Geotecnica, San José, Costa Rica Marco Avanzini Museo Tridentino di Scienze Naturali, Trento, Italy David Berman Carnegie Museum of Natural History, Pittsburgh, PA, USA Brent Breithaupt Laramie, WY, USA William DiMichele National Museum of Natural History, Washington, D.C., USA John R. Foster Museum of Western Colorado, Grand Junction, CO, USA Gerard Gierlinski Polish Geological Institute, Warsaw, Poland Jean Guex University of Lausanne, Lausanne, Switzerland Jerald D. Harris Dixie State College, St. George, UT, USA Andrew B. Heckert Appalachian State University, Boone, NC, USA Adrian P. Hunt Flying Heritage Collection, Everett, WA, USA Hendrik Klein Neumarkt, Germany Karl Krainer University of Innsbruck, Innsbruck, Austria Martin G. Lockley University of Colorado at Denver, Denver, CO, USA Claudia Marsicano Universidad de Buenos Aires, Buenos Aires, Argentina Gary S. Morgan NMMNHS Donald R. Prothero Occidental College, Los Angeles, CA, USA Silvio Renesto Università degli Studi dell’Insubria, Varese, Italy Joerg W. Schneider Technical University BergAkademie of Freiberg, Freiberg, Germany Jingeng Sha Nanjing Institute of Geology and Palaeontology, Nanjing, China Sebastian Voigt Urweltmuseum GEOSKOP/Burg Lichtenburg, Thallichtenberg, Germany Ralf Werneburg Naturhistorisches Museum Schloss Bertholdsburg, Schleusingen, Germany Richard S. White, Jr. International Wildlife Museum, Tucson, AZ, USA NEW MEXICO MUSEUM OF NATURAL HISTORY AND SCIENCE BULLETINS

27. Late Triassic microvertebrates from the lower Chinle Group (Otischalkian-Adamanian: Carnian), 2004. by Andrew B. Heckert, 170 pp. 28. New Mexico’s Ice Ages, 2005. edited by Spencer G. Lucas, Gary S. Morgan, and Kate E. Zeigler, 300+ pp. 29. Vertebrate Paleontology in Arizona, 2005. edited by Andrew B. Heckert & Spencer G. Lucas, 210 pp. 30. The Nonmarine Permian, 2005. edited by Spencer G. Lucas & Kate E. Zeigler, 362 pp. 31. Permian of Central New Mexico, 2005. edited by Spencer G. Lucas, Kate E. Zeigler & Justin A Spielmann, 176 pp. 32. Genética y Mamíferos Mexicanos: Presente y Futuro, 2006. edited by Ella Vázquez-Domínguez and David J. Hafner, 73 pp. 33. Skeletal Impact of Disease, 2006. by Bruce M. Rothschild and Larry D. Martin, 226 pp. 34. America’s Antiquities: 100 Years of Managing Fossils on Federal Lands, 2006. edited by Spencer G. Lucas, Justin A. Spielmann, Patricia M. Hester, Jason P. Kenworthy and Vincent L. Santucci, 185 pp. 35. Late Cretaceous Vertebrates from the Western Interior, 2006. edited by Spencer G. Lucas and Robert M. Sullivan, 410 pp. 36. Paleontology and Geology of the Upper Jurassic Morrison Formation, 2006. edited by John R. Foster and Spencer G. Lucas, 249 pp. 37. The Triassic-Jurassic Terrestrial Transition, 2006. edited by Jerry D. Harris, Spencer G. Lucas, Justin A. Spielmann, Martin G. Lockley, Andrew R.C. Milner and James I. Kirkland, 607 pp. 38. Pennsylvanian-Permian Fusulinaceans of the Big Hatchet Mountains, New Mexico, 2006. by Garner L. Wilde, 331 pp. 39. Upper Aptian-Albian Bivalves of Texas and Sonora: Biostratigraphic, Paleoecologic and Biogeographic Implications, 2007. edited by Robert W. Scott, 39 pp. 40. Triassic of the American West, 2007. edited by Spencer G. Lucas and Justin A. Spielmann, 247 pp. 41. The Global Triassic, 2007. edited by Spencer G. Lucas and Justin A. Spielmann, 415 pp. 42. Cenozoic Vertebrate Tracks and Traces, 2007. edited by Spencer G. Lucas, Justin A. Spielmann and Martin G. Lockley, 330 pp. 43. The Late Triassic archosauromorph Trilophosaurus, 2008. by Justin A. Spielmann, Spencer G. Lucas, Larry F. Rinehart and Andrew B. Heckert, 177 pp. 44. Neogene Mammals, 2008. edited by Spencer G. Lucas, Gary S. Morgan, Justin A. Spielmann and Donald R. Prothero, 442 pp. 45. The Paleobiology of Coelophysis bauri (Cope) from the Upper Triassic (Apachean) Whitaker quarry, New Mexico, with detailed analysis of a single quarry block, 2009. by Larry F. Rinehart, Spencer G. Lucas, Andrew B. Heckert, Justin A. Spielmann and Matthew D. Celeskey, 260 pp. 46. The taxonomy and paleobiology of the Late Triassic (Carnian-Norian: Adamanian-Apachean) drepanosaurs (Diapsida: Archosauromorpha: Drepanosauromorpha, 2010. by Silvio Renesto, Justin A. Spielmann, Spencer G. Lucas and Giorgio Tarditi Spagnoli, 81 pp. 47. Ichnology of the Upper Triassic (Apachean) Redonda Formation, east-central New Mexico, 2010. by Spencer G. Lucas, Justin A. Spielmann, Hendrik Klein and Allan J Lerner, 75 pp. 48. New Smithian (Early Triassic) ammonoids from Crittenden Springs, Elko County, Nevada: Implications for taxonomy, biostratigraphy and biogeography, 2010. by James F. Jenks, Arnaud Brayard, Thomas Brühwiler and Hugo Bucher, 41 pp. 49. Carboniferous-Permian transition in Cañon del Cobre, northern New Mexico, 2010. edited by Spencer G. Lucas, Jörg W. Schneider and Justin A. Spielmann, 229 pp. 50. Review of the tetrapod ichnofauna of the Moenkopi Formation/Group (Early-Middle Triassic) of the American Southwest, 2010. by Hendrik Klein and Spencer G. Lucas, 67 pp. 51. Crocodyle tracks and traces, 2010. edited by Jesper Milàn, Spencer G. Lucas, Martin G. Lockley and Justin A. Spielmann, 244 pp. 52. Selachians from the Upper Cretaceous (Santonian) Hosta Tongue of the Point Lookout Sandstone, central New Mexico, 2011. by Jim Bourdon, Keith Wright, Spencer G. Lucas, Justin A. Spielmann and Randy Pence, 54 pp. 53. Fossil Record 3, 2011. edited by Robert M. Sullivan, Spencer G. Lucas and Justin A. Spielmann, 736 pp. 54. Ichnology of the Mississippian Mauch Chunk Formation, eastern Pennsylvania, 2012. by David L. Fillmore, Spencer G. Lucas and Edward L. Simpson, 136 pp. 55. Tetrapod fauna of the Upper Triassic Redonda Formation, east-central New Mexico: The characteristic assemblage of the Apachean land-vertebrate faunachron, 2012. by Justin A. Spielmann and Spencer G. Lucas, 119 pp. 56. Revision of the Lower Triassic tetrapod ichnofauna from Wióry, Holy Cross Mountains, Poland, 2012. by Hendrik Klein and Grzegorz Niedzwiedzki, 62 pp. 57. Vertebrate Coprolites, 2012. edited by Adrian P. Hunt, Jesper Milàn, Spencer G. Lucas and Justin A. Spielmann, 387 pp. 58. A new archaic basking shark (Lamniformes: Cetorhinidae) from the late Eocene of western Oregon, U.S.A., and description of the dentition, gill rakers and vertebrae of the recent basking shark Cetorhinus maximus (Gunnerus), 2013. by Bruce J. Welton, 48 pp. 59. The Carboniferous-Permian transition in central New Mexico, 2013. edited by Spencer G. Lucas, W. John Nelson, William A. DiMichele, Justin A. Spielmann, Karl Krainer, James E. Barrick, Scott Elrick and Sebastian Voigt, 389 pp. 60. The Carboniferous-Permian transition, 2013. edited by Spencer G. Lucas, William A. DiMichele, James E. Barrick, Joerg W. Schneider and Justin A. Spielmann, 465 pp. 61. The Triassic System: New Developments in Stratigraphy and Paleontology, 2013. edited by Lawrence H. Tanner, Justin A. Spielmann and Spencer G. Lucas, 612 pp. 62. Fossil Footprints of Western North America, 2014. edited by Martin G. Lockey and Spencer G. Lucas, 508 pp. 63. Variation in the Dentition of Coelophysis bauri, 2014. by Lisa G. Buckley and Philip J. Currie, 73 pp. Michael J. Orchard, 2014, Conodonts from the Carnian-Norian Boundary (Upper Triassic) of Black Bear Ridge. New Mexico Museum of Natural History and Science Bulletin 64. CONODONTS FROM THE CARNIAN-NORIAN BOUNDARY (UPPER TRIASSIC) OF BLACK BEAR RIDGE, NORTHEASTERN BRITISH COLUMBIA, CANADA

MICHAEL J. ORCHARD

Contents Abstract 1 INTRODUCTION 2 Previous Work 2 THE CONODONT FAUNAS 3 Taxonomic method and generic identity 8 Evolutionary trajectories 8 Quadralella 8 Kraussodontus 10 Metapolygnathus 10 Parapetella 10 Carnepigondolella 20 Acuminatella 20 Primatella 25 Epigondolella 25 CONODONT ZONATION 25 Carnepigondolella samueli Zone 26 Carnepigondolella eozoae-K. ludingtonensis Subzone 26 Carnepigondolella zoae Subzone 27 Carnepigondolella medioconstricta Subzone 29 Carnepigondolella spenceri Subzone 30 Primatella primitia Zone 30 Acuminatella sagittale-Parapetella beattyi Subzone 31 Acuminatella angusta-Metapolygnathus dylani Subzone 31 Acuminatella acuminata-Parapetella prominens Subzone 32 Metapolygnathus parvus Subzone 32 Lower subdivision 32 Middle subdivision 32 Upper subdivision 33 Primatella asymmetrica-Norigondolella sp. Subzone 33 Epigondolella quadrata Zone 33 Epigondolella triangularis Zone 33 INTEGRATED BIOSTRATIGRAPHY AT BLACK BEAR RIDGE 33 Complementary data for the CNB placement in British Columbia 34 Correlation with Pizzo Mondello, Sicily 38 Conodont datums for the CNB 40 SUMMARY 40 ACKNOWLEDGMENTS 41 SYSTEMATIC PALEONTOLOGY 41 Foreword 41 Acuminatella 41 Carnepigondolella 47 Epigondolella 55 Kraussodontus 59 Metapolygnathus 66 Norigondolella 71 Parapetella 72 Primatella 87 Quadralella 106 Misikella 127 REFERENCES 127 APPENDIX I 130 Tables 1-8 APPENDIX II 138 List of conodont taxa View of Peace Reach, Williston Lake looking north toward Black Bear Ridge. Michael J. Orchard, 2014, Conodonts from the Carnian-Norian Boundary (Upper Triassic) of Black Bear Ridge. New Mexico Museum of Natural History and Science Bulletin 64. 1 CONODONTS FROM THE CARNIAN-NORIAN BOUNDARY (UPPER TRIASSIC) OF BLACK BEAR RIDGE, NORTHEASTERN BRITISH COLUMBIA, CANADA

MICHAEL J. ORCHARD Natural Resources Canada, Geological Survey of Canada, 1500-605 Robson Street, Vancouver, BC, V6B 5J3. Canada. Email: [email protected]

Abstract—Conodonts from the Carnian-Norian Boundary (CNB) interval at the Global Stratigraphic Section and Point (GSSP) candidate section at Black Bear Ridge (BBR), British Columbia, Canada, include five previously introduced genera (Acuminatella, Kraussodontus, Parapetella, Primatella, Quadralella), 71 new species (plus nine previously named, and 13 more in open nomenclature), and 47 new morphotypes. These elements, and others previously known, display progressive morphogenesis through the strata of the Ludington and Pardonet formations, which represent a continuous Upper Triassic slope-basin succession at the western edge of Pangea. Two conodont zones and nine subzones are defined for the interval, in ascending stratigraphic order:Carnepigondolella samueli Zone with the subzones of C. eozoae-Kraussodontus ludingtonensis, C. zoae, C. medioconstricta, and C. spenceri; the Primatella primitia Zone with the subzones of Acuminatella sagittale-Parapetella beattyi, A. angusta-Metapolygnathus dylani, A. acuminata- Pa. prominens, M. parvus (three subdivisions), and Primatella asymmetrica-Norigondolella sp. These strata are capped by the Early Norian Epigondolella quadrata Zone. Conodont faunal turnovers are identified at the boundary between the samueli and primitia zones with the extinction of Carnepigondolella, and between the primitia and quadrata zones with the extinction of Acuminatella and Primatella. However, the most significant turnover is in the boundary interval of theacuminata -prominens and parvus subzones where, respectively, 16 and 46 taxa disappear, including all Quadralella, Kraussodontus, and most Parapetella species. This turnover falls within a 5 m boundary interval bracketed by diagnostic ammonoids of the Upper Carnian Klamathites macrolobatus Zone and those of the Lower Norian Stikinoceras kerri Zone. The undated boundary interval includes an organic carbon isotope minimum, and the first occurrences of the bivalve Halobia austriaca, and the ammonoid Pterosirenites. Conodonts from this boundary interval are also identified in association with H. austriaca and Pterosirenites at nearby Pardonet Hill east (PHE). Matrix from archival ammonoid collections of the Macrolobatus Zone has also yielded the boundary conodont faunas, implying that the entire boundary interval at BBR is equivalent to the Macrolobatus Zone. Broad correlation with the GSSP candidate section at Pizzo Mondello (PM) in Sicily can be achieved at several levels around the parvus Subzone: at its base with the appearance of the name-giver; at levels within it based on the appearances of Parapetella destinae, Parapetella irwini, and Primatella bifida; and at its top where typical Carnian taxa disappear (equivalent to event T3 at PM). At BBR, the parvus Subzone documents the decline and extinction of Carnian conodont genera, the bloom of diminutive taxa, and the ascendency of the Primatella stock, precursor of Epigondolella. The parvus Subzone includes significant conodont, ammonoid, and bivalve appearances that could serve as indices or proxies for the CNB, but use of any of them would have the effect of assigning some Macrolobatus Zone collections to the Norian Stage. The base of the asymmetrica-Norigondolella Subzone of the primitia Zone offers a CNB position that is most closely aligned with the traditional base of the Norian, i.e. at the base of the Kerri Zone, and only that position assigns all Anatropites-bearing collections to the Carnian. At BBR, this level is defined by the disappearance of Metapolygnathus parvus and its diminutive associates rather than by new appearances. Uncommon taxa that appear at that time may be endemics: Acuminatella curvata at BBR, and Primatella? gulloae at PM. The disappearance of M. parvus and its associates close the Carnian chapter in conodont evolution, and their extinction may be favored as a natural boundary. Such a position for the CNB does, however, remove Halobia species as definitive indices, and places the range of Pterosirenites on both sides of the Carnian-Norian Boundary. Subsequent floods of Norigondolella, initially N. norica, provide a strong and definitive Norian signal at BBR and nearby sections. In addition to five new genera and nine new species previously introduced from BBR, the following new taxa are named: Acuminatella binodosa, A. constricta, A. curvata, A. denticulata, A. longicarinata, A. sagittale, A. sinuosa, A.? extensa, A.? prima, Carnepigondolella anitae, C. gibsoni, C. milanae, C. postsamueli, C. spenceri, Kraussodontus ludingtonensis, K. margaretae, K. roberti, K. rosiae, K. urbanae, K. vancouverense, K. wendae, Metapolygnathus dylani, Norigondolella norica, Parapetella beattyi, P. broatchae, P. clareae, P. columbiense, P. cordillerense, P. destinae, P. elegantula, P. hillarae, P. irwini, P. johnpauli, P. lanei, P. posterolata, P. prominens angulare, P. p. circulare, P. pumilio, P. riteri, P. rubae, P. willifordi, Primatella bifida, P. circulare, P. elongata, P. mclearni, P. oblonga, P. ovale, P. posteroglobosa, P. rectangulare, P. rhomboidale, P. rotunda, P. stanleyi, P. subquadrata, P. triangulare, P. vanlierae, Quadralella deflecta, Q. karenae, Q. kathleenae, Q. mcrobertsi, Q. pardoneti, Q. posteroexpansa, Q. postlobata, Q. praecommunisti curvata, Q. p. ornata, Q. roysi, Q. sigmoidale, and Q. willistonense. 2 INTRODUCTION Black Bear Ridge (BBR) in northeastern British Columbia, The succession of conodonts recovered from strata at BBR reveals Canada (Fig. 1) is an exceptional locality for richly fossiliferous progressive morphogenesis in multiple Upper Carnian clades, far more Upper Triassic strata. As such, it has an important role to play in the than previously identified. These provide numerous datums for potential development of the Triassic part of the International Time Scale. The correlation and definition. Based on a substantially new taxonomy, International Commission on Stratigraphy recognizes the locality as a evolutionary trajectories of seven genera, five of them new (Orchard, leading candidate for definition of the Global Stratigraphic Section and 2013), are traced through the boundary interval. A total of 118 conodont Point (GSSP) for the base of the Norian Stage of the Upper Triassic species and subspecies, and 47 morphotypes form the basis of new Series (Ogg, 2012, p. 719). On a national scale, Upper Triassic rocks of conodont zones and subzones that are integrated with other fossil and northeastern British Columbia are a significant hydrocarbon resource in geochemical data from the section to produce a temporal framework for the Western Canadian Sedimentary Basin, and the fossil tools described the GSSP candidate, as well as for application in the Western Canada here are crucial in that basin analysis. Sedimentary Basin, and demonstrably beyond (Carter and Orchard, Black Bear Ridge and other localities in the former Peace River 2013; Balini et al., 2014). Valley yielded outstanding ammonoid faunas that were foundational for Previous Work the standard North American Upper Triassic chronology (Tozer, 1967, 1994). More recently, conodont faunas recovered in direct association Black Bear Ridge and other important sections crop out on the east- with the ammonoids, and from intervening strata, were intercalibrated west arm of Williston Lake, which was created when the Peace River with these ammonoid zones (Orchard, 1983, 1991b). Realization that valley was flooded after the completion of the W.A.C. Bennett Dam in BBR provided a remarkable conodont record around the Carnian- 1968. Due to seasonal rise and fall of water levels, considerable new Norian boundary (CNB) led to this systematic study of the conodonts outcrop appeared around the perimeter of the lake during the following and their role in correlation and chronostratigraphic definition. decade. New successions of richly fossiliferous Upper Triassic strata The BBR section occurs on the northern shore of Peace Reach on were discovered during reconnaissance fieldwork by E.T. Tozer and Williston Lake (Fig. 1). The study interval reported here comprises the the present author in 1980 and 1981. Prior to this, both McLearn lower 90 m of the section, which embraces all options for the definition (1947, 1960) and Tozer (1965, 1967) had worked in the former Peace of the CNB. The lower half of the section is assigned to the Ludington River Valley to produce a North American Upper Triassic ammonoid Formation, and the upper part to the Pardonet Formation (Fig. 2). At chronology (Silberling and Tozer, 1968). The flooding inundated many this locality, it is believed that an essentially complete record of CNB classic Triassic sites but also exposed fresh outcrops from which large strata exists, representing a deep-water slope and basin setting at the collections of macrofossils and conodont samples were made. western edge of Pangea (Zonneveld et al., 2010a, b). Black Bear Ridge Much of the Peace River ammonoid data were incorporated in lies west of a hinge line that separated medial to distal ramp facies of the authoritative account of Triassic Ammonoidea provided by Tozer the Ludington Formation from localities farther east where equivalent (1994). Abundant conodonts recovered as a result of the collaborative strata are represented by the proximal- to medial-ramp strata of the fieldwork resulted in a tightly integrated ammonoid-conodont Baldonnel Formation (Zonneveld and Orchard, 2002). The Pardonet biozonation from the Late Carnian Macrolobatus Zone through the Formation, which represents a distal ramp facies, transgressed eastward Upper Norian Cordilleranus Zone (Orchard, 1983, 1991b). The and has a diachronous base. At BBR, the CNB lies within the Pardonet importance of the BBR section for the CNB was realized after several Formation, but in eastern outcrops it occurs within the shallow-water visits to the lake and increasingly more detailed studies that culminated Baldonnel Formation. in bed-by-bed collections from the key intervals of faunal change.

Figure 1. Location of Black Bear Ridge (5) on Peace Reach, northeastern British Columbia. Other key sections are numbered as follows: 1. Carbon Creek, 2. McClay Spur, 3. Glacier Spur, 4. Brown Hill, 6. Pardonet Hill east, 7. Juvavites Cove. Inset shows location of study area, and also of Haida Gwaii, in British Columbia, western Canada. 3

Figure 2. Photograph of lower outcrop on Black Bear Ridge, showing transition from Ludington to Pardonet formations. The gash in the center of the photo marks slight bedding parallel fault movement within the lower Kerri Zone (beds 25-26): the contrasting appearance results from differing weathering styles. Circled is a human scale, located in the Dawsoni Zone. The asterisk marks the Carnian-Norian boundary interval. Black Bear Ridge was first visited by the author in 1981. The Orchard (2013) introduced five new conodont genera and provided immediate highlight was an excellent section with abundant ammonoids a preliminary summary of the biostratigraphy and zonation of the and bivalves that indicated Middle Norian (Columbianus Zone) up upper Ludington and lower Pardonet formations at BBR (Fig. 4). The to Hettangian (Lower Jurassic) strata. This part of the outcrop was application of the new zonal conodont scheme has subsequently been sampled for conodonts in moderate detail, but only isolated samples demonstrated on both Haida Gwaii (Carter and Orchard, 2013), and in were taken from the lower section, where uncommon indicators of the Nevada (Balini et al., 2014). Lower Norian Kerri and Dawsoni ammonoid zones were identified. Two collections of conodonts were recovered initially, and assigned to THE CONODONT FAUNAS the “Paragondolella polygnathiformis” and “Epigondolella primitia” A minimum of 35,000 conodont elements, extracted from over zones, or zones 19 and 20 of Sweet et al. (1971), broad intervals that 230 kg of carbonate rock from the lower BBR section, were examined were the sum of the zonation at that time. The first collection included for this study (Appendix, Tables 1, 2). Samples collected in 2004 are ornate forms that were then included in “Epigondolella abneptis”, but a not included in these totals, nor are the many thousands of elements decade later were separated as Carnepigondolella samueli. recovered from contemporaneous strata, particularly from Pardonet Black Bear Ridge was re-visited in 1982 with the primary focus Hill east (PHE) and nearby at Juvavites Cove (Fig. 1). In this work, on the upper part of the formation and the Triassic-Jurassic boundary 1638 individual Scanning Electron Microphotographs of 570 featured (Tozer, 1982; Sephton et al., 2002; Wignall et al., 2003, 2007; Hall P1 conodont elements are illustrated, in most cases including three and Pitaru, 2004). Additional samples were taken from the sub- views (upper, lower, lateral). This lavish illustration is exceptional Columbianus Zone section of the Pardonet Formation, but the lower documentation for a conodont fauna and provides the taxonomic part of the section was first sampled systematically, at ~3 m intervals, in foundation for the conclusions reached herein. 1992. The results established the overall biostratigraphic record for the The conodonts are generally well preserved, abundant, and often entire BBR section (Orchard et al., 2001b, c). A farther decade passed morphologically similar yet very variable. The plasticity of Late before a more detailed, multidisciplinary project could begin on the Triassic conodonts has often hampered their study, but what becomes CNB interval. clearer with closer examination aided by scanning electron microscope In 2001-02, samples were collected at ~1 m intervals through the (SEM) imagery is a remarkable record of evolutionary morphogenesis lower ~20 m of the Pardonet Formation, and from ~2 m intervals for through the Upper Carnian Macrolobatus zone of the standard a further ~20 m, ending near the top of the Kerri Zone. Sampling was ammonoid chronology. The conodonts are generally dark brown in also extended downward through the upper 30 m of exposed Ludington colour, expressed as a Colour Alteration Index (CAI) of 3.5-4. The best Formation (Fig. 3). In 2004, the balance of the exposed Ludington preserved elements were from the cleaner carbonates, whereas those Formation was sampled over ~20 m, and key intervals of faunal change from the shaly interbeds were often partly obscured by adhering matrix. identified near the base of the Pardonet Formation were sampled Abundance was variable, but few samples were barren. Conodont yields bed-by-bed. Subsequently, focus shifted to the CNB interval as now ranged from a few, to over a thousand elements per kilogram, and were recognized and, in 2007, there followed intensive sampling for both commonly between a few tens to a few hundred elements /kg. (Fig. 5). micro- and macrofossils (Orchard, 2007b, c, 2010b, 2011). As part of The most abundant collections came from a ~4 m CNB interval where this multidisciplinary effort, the sedimentary framework of the section a faunal turnover involves the disappearance of 63 taxa belonging to was studied by Zonneveld et al. (2010a, b), its bivalves by McRoberts four genera, and the appearance of 33 new taxa largely in the lower (2011), the ammonoids by Krystyn and Balini (Balini et al., 2012), and 2 m, and many of them short-lived (Fig. 6). Yield drops dramatically organic carbon isotope chemostratigraphy by Williford et al. (2007). mid-way through this interval, as it does after two other abundance These results confirmed that a narrow CNB interval bracketed by peaks, at the top of the samueli and primitia zones. Each of these three Carnian ammonoids below and by those of Norian age above embraced peaks corresponds to a faunal turnover that involved the extinction and faunal changes in conodonts and bivalves, and also corresponded to a appearance of both species and genera (Fig. 5). Details of conodont C isotope minimum. Supplementary sampling was undertaken in 2010 element numbers and yields provided in Tables 1 and 2 (Appendix) to examine in detail the succession between the primitia and quadrata distinguish platform and ramiform elements. The latter are severely conodont zones at the top of the sampled section. under-represented in the collections and invariably fragmentary, so are Employing the substantially revised taxonomy provided herein, not considered further here. 4

Figure 3. Columnar section of Black Bear Ridge lower section, adapted from Zonneveld et al. (2010), to which the reader is referred for a discussion of the sedimentary units on the left. The central columns show successive years of conodont sampling with bed/ sample numbers indicated. The occurrence of macrofossils, and of diagnostic ammonoid genera are shown: A=Anatropites, G=Guembelites, S=Stikinoceras, and M=Malayites: these constrain the ammonoid zones shown to the right of the stratigraphic column; Dw = Dawsoni Zone. Also shown are samples with common Norigondolella, indicated by a black dot. The new conodont zonation is shown on the right. 5

Figure 4. Conodont zonation across the Carnian-Norian Boundary at Black Bear Ridge and its evolution during the last 30 years as a result of an increasingly refined taxonomy: oldest on left, latest on right. See text for details. The stratigraphic column and data to its left as in Fig. 3, and the bed numbers adopted as standard are shown to its right. Conodont generic abbreviations are Ca. =Carnepigondolella, K. = Kraussodontus, Me. =Metapolygnathus, Ac. =Acuminatella, Pa. =Parapetella, Pr. =Primatella, Ep. =Epigondolella. 6

Figure 5. Conodont yield through the Black Bear Ridge section, ranges of conodont genera, and the suggested position of the faunal turnovers T1-T3 identified by Mazza et al. (2010) at Pizzo Mondello. Columnar section on left as in Fig. 3, conodont zones on right as in Fig. 4. Note the yields are highest around the zonal boundaries and in the CNB boundary interval around the middle of the primitia Zone: these levels also coincide with generic turnovers. The largest disappearance of species is in the CNB boundary interval where yields reach 1000+ /kg. See Fig. 6 for details. 7

Figure 6. Bed-by-bed analysis of conodont fauna for a 5 m boundary interval in the Pardonet Formation between the highest Macrolobatus (Anatropites) and lowest Kerri (Guembelites) Zone ammonoid indicators. Generally high yields associated with diverse, typically Carnian conodonts (pale gray bars) drop off as that fauna disappears and diminutive derivatives (black bars) bloom prior to their extinction, after which the fauna is almost exclusively Primatella and Acuminatella (medium gray bars). Norigondolella blooms appear later, within the Kerri Zone. Indicated bed/ sample numbers are shown next to total taxa present, the number of last appearances (LAD), and first appearances (FAD). The appearance of the first diminutive Parapetella in bed 18f coincides with the last dominant Carnian fauna, and corresponds to the peak negative organic carbon isotopes perturbation reported by Williford et al. (2007), indicated by the arrow on right. This also aligns with the appearance of Halobia austriaca and a proposed position for the CNB (McRoberts and Krystyn, 2011): see Figure 28. 8 Taxonomic Method and Generic Identity elevated, generally non-nodose anterior platform margins. Finally, Students of Upper Triassic conodonts have traditionally adopted a Metapolygnathus is now restricted to a lineage characterized by mostly classification that emphasizes few genera, and constituent species that inornate elements with relatively flat platform margins, and medial to display a wide range of intraspecific morphological variability. Broad anterior pits. Many of the Upper Carnian genera exhibit an evolutionary interpretations of species has arisen from inadequate illustration and trend that involves anterior migration of the pit, and so an anterior pit is description of taxa introduced half a century ago, in some cases with no longer a sole defining criterion forMetapolygnathus . little or no stratigraphic context, as in the Hayashi (1968a, b) mixed Both in Metapolygnathus, and in some lineages of fauna. The present work adopts a strict typological approach, with Carnepigondolella and Parapetella, a trend of progressive diminution species defined narrowly, and additional subspecies or morphotypes in platform size, and concomitant lengthening of the free blade is differentiated to capture variation that might otherwise be concealed. In observed through time. In Carnepigondolella this reaches its acme this way, the stratigraphic utility of the BBR collections is optimized. near the top of the samueli Zone, whereas in the other two genera these Future recognition of different spectra of species and morphotypes trends reach a maximum in the parvus Subzone of the CNB interval. elsewhere may improve our understanding of both temporal and This diminution precedes the extinction of clades of Carnepigondolella biogeographic differences. and, later, of Parapetella. The classification presented here is essentially a phenetic In addition to the platform-bearing genera described above, taxonomy: conodont P1 platform elements have been organized Norigondolella becomes common only above the CNB interval at according to their morphological similarities as determined by empirical BBR, although rare specimens occur below and within the boundary observation. Resulting groups of similar elements have been considered interval. Successive faunal blooms of two Norigondolella species in the context of their stratigraphic occurrence within the BBR section. occur in undisputed Norian strata, presumably reflecting changing This led to proposed conodont phylogenies that incorporate plausible environmental conditions, and possibly related to ocean cooling (Rigo trajectories of morphological changes through successive Late Carnian- et al., 2014). Finally, single elements of Misikella occur in two samples, Early Norian faunas. In many cases, similar evolutionary trends can be and are included here for the sake of completion. recognized in successive or parallel lineages, e.g., anterior migration Evolutionary Trajectories of the pit, platform shortening/ blade lengthening, anterior margin differentiation. Iterative evolution results in similar characteristics This part presents hypotheses on the morphogenesis of most emerging separately in Carnepigondolella, Metapolygnathus, and common platform elements of the CNB section at BBR. Proposed Parapetella, and to a lesser extent in Acuminatella and Primatella. phyletic linkages are described and illustrated in 19 figures (Figs. With the introduction of five new conodont platform genera, the 7-25). The generally inornate genera are described first, followed by number of genera recognized in the study interval is doubled. In addition the highly ornate genera. to 27 known species, BBR has yielded 71 new species (plus 13 more Quadralella (Figs. 7-10; 75-88) in open nomenclature), six new subspecies, and 47 new morphotypes. In support of this taxonomy, numerous elements are illustrated in three The majority of Late Carnian taxa at BBR are assigned to this standard views and orientations, and at uniform magnification. With genus, as are many Early Carnian taxa derived from Gondolella this monographic treatment of boundary interval conodonts, it is hoped polygnathiformis Budurov and Stefanov sensu lato, the first that future comparisons of contemporary faunas and their correlation representative of Quadralella. It encompasses several species will be less ambiguous. formerly included in “Metapolygnathus nodosus” by this author, or Recent literature on Upper Triassic conodonts includes platform in Paragondolella species by others. P1 elements of Quadralella are elements assigned to species of Carnepigondolella, Epigondolella, often sub-rectangular in outline but variation in their posterior platform Metapolygnathus, Norigondolella, and Paragondolella. The last of development provides a basis for speciation. All species have relatively these, based on Middle Triassic Paragondolella excelsa, is regarded as poorly differentiated anterior platform nodes or none at all. Growth an inappropriate genus for most Upper Triassic species (Orchard, 2013). series of some species are uncertain, and some taxa are only recognized Paragondolella elements differ in having relatively flat platforms on the basis of “mature” individuals. and very high carinas, as well as a different multielement apparatus Representatives of Quadralella are extant from the base of the (Orchard, 2005). The assertion by Kiliç et al. (2013, p. 64) that the BBR section, but most species disappear within the parvus Subzone. S0 element of Paragondolella identified by Orchard (2005, figs. 20G Adults of the oldest species fall into two subgroups: those with lateral and 21G) is identical to the S3 elements of Neogondolella (Orchard, platform constrictions, as exemplified by Q. lobata Orchard, and those 2005, fig. 10E) is incorrect. The former element has two bilaterally that are more uniformly rectangular in outline, as in Q. carpathica symmetrical antero-lateral processes, whereas the latter has a markedly (Mock). Early growth stages of all these taxa (included in Q. spp. asymmetrical anterior part with a short antero-lateral process rising indet.) appear to have a more developed anterior platform and a reduced from the anterior process. posterior platform, but they cannot be unequivocally assigned to Epigondolella was initially a hold-all for all Late Triassic ornate particular ‘mature’ species. elements and continues to be used for some Late Carnian species. In Quadralella lobata has a very short anterior free blade but a this work the genus is only used for Norian species of E. quadrata progressive reduction in the lateral flanges on the margin of the blade and its associates. The species content of Norigondolella is generally as well as in the length of the platform, results in a more prominent free undisputed nowadays, although both Gondolella and Neogondolella blade in the younger Q. postlobata sp. nov. (Fig. 7). The cusp in the have been used in the literature for those species. With these taxonomic center of the constricted posterior platform is a feature of both species, constraints in mind, and the knowledge that pit position is very variable and of the probable derivative Q. karenae sp. nov., in which the posterior in Carnian taxa, Orchard (1991a, b) assigned most other Carnian platform beyond the constriction is strongly reduced. The younger species to Metapolygnathus pending a clearer understanding of phyletic Quadralella sp. nov. A has a very similar outline to Q. postlobata but relationships. The introduction by Kozur (2003) of Carnepigondolella has more strongly developed anterior nodes, although much less so than for one Carnian lineage (Orchard, 1991a, fig. 4) was a useful first the slightly younger homeomorph Primatella circulare sp. nov. revision, but even that genus has been very broadly interpreted recently. Longer and narrower platform elements that have a carina and In a preliminary report of this work (Orchard, 2013), five new cusp similar to that of Quadralella postlobata but with a less expanded conodont genera were introduced as further rationalization of Upper posterior platform, and a medial constriction rather than a posterior Triassic conodont taxonomy. Two new ornate genera, Acuminatella one, are assigned to Q. pardoneti sp. nov. (Fig. 7). Two morphotypes and Primatella, were differentiated from elements formerly assigned of this species are distinguished on the basis of their pit position. It to the broad Epigondolella primitia population of Orchard (1983), and is possible that these two morphotypes are early growth stages of, later to Metapolygnathus primitius (Orchard, 1991a, b). Acuminatella respectively, Q. willistonensis sp. nov. and Q. praecommunisti (Mazza, accommodates those taxa with pointed platforms that Kozur (2003) Rigo and Nicora), although their ranges are not identical. The latter tentatively assigned to Orchardella. Quadralella encompasses many two species (Fig. 8) are based on relatively large specimens that have of the inornate or weakly ornate, subrectangular platform elements neither a posterior constriction nor, usually, a terminal cusp. Other, formerly included by many authors in Paragondolella, or by this partly contemporaneous elements similar to Q. pardoneti differ in their author in Metapolygnathus. Kraussodontus embraces forms with long carinas, their sometimes very narrow posterior platforms, and relatively narrow and elongate, roundly terminated platforms. A their invariably posterior pits. For the purposes of this report, these fifth genus, Parapetella, was introduced for elements that develop small elements are separated as Quadralella spp. indet., alpha, beta, and gamma morphotypes (Fig. 7). They probably represent juveniles of 9

Figure 7. Proposed phyletic linkages in small- to medium-sized elements of Quadralella that have a lobate or reduced platform posterior of a constriction; free blade lengthens and position of constriction changes through time. The Black Bear Ridge columnar section is depicted on the left with bed numbers. Solid dots are occurrences in beds indicated and open circles are uncertain occurrences. Vertical lines: solid = observed ranges, dashed = uncertain ranges. Horizontal or diagonal dashed lines are proposed lines of derivation; dotted lines are more tenuous linkages, or connect related but stratigraphically separated elements. 10 the aforementioned taxa but they have a longer range and may include have lateral indentations on both margins rather than the one seen in early growth stages of additional species, e.g., Q. angulata (Mazza, Cau K. aff. margaretae; they differ from each other in platform length and and Rigo) and Q. noah (Hayashi). posterior platform outline. Rare K. sp. nov. A is also similar to K. aff. Several species of Quadralella are defined on relatively large margaretae but has a relatively reduced platform. platform elements in which diagnostic criteria are well displayed. Relatively narrow Kraussodontus may have arisen from the These larger elements include the relatively narrow Q. angulata (Fig. broader Quadralella prior to the stratigraphic record preserved at Black 8) and the broader Q. carpathica (Fig. 9), which have a short free blade, Bear Ridge. In turn, the genus may have been precursor to the pointed and their proposed younger derivatives Q. tuvalica (Mazza and Rigo) platforms manifest in some Acuminatella and Parapetella species, and Q. willistonensis sp. nov., which have distinctive free blades. All the first through anterior node differentiation, and the latter through these species have a posterior pit near the end of the keel, as does Q. anterior parapet growth. kathleenae sp. nov. (Fig. 8), which differs in its more developed anterior platform nodes. Anterior shifting of the pit is a trend that begins in Q. Metapolygnathus (Figs. 13; 46-48) noah and culminates in Q. praecommunisti, in which a keel extends This present author chose in the past to assign most Upper Carnian well posterior of the submedial pit. Mazza et al. (2011) suggested that taxa to this genus pending a thorough taxonomic study, which this this lineage led to Metapolygnathus communisti Hayashi, but similar work purports to be. Metapolygnathus is now restricted to generally elements (M. ex gr. communisti of this report) were extant earlier than subrectangular, unornamented elements lacking parapets and with pits Q. praecommunisti. In this work, the latter species is subdivided into located in a medial or anterior platform position (Fig. 13). Unlike many three subspecies. Tethyan regions, the group of M. communisti is rare at BBR, and those An uncommon species that appears before Q. praecommunisti is that occur are sufficiently variable to differentiate seven morphotypes Q. mcrobertsi sp. nov. This species, like Q. kathleenae, has relatively that succeed each other through the section. Notably, none are identical well developed anterior nodes, but it differs in having a submedial pit to the holotype from Japan. and may have been an early offshoot from Q. noah. This entire group The Metapolygnathus ex gr. communisti elements show various of subrectangular elements (Fig. 8) has a complex history that lacks trends, one involving increased elevation of the anterior platform an obvious juvenile dimension, although that is assumed to lie in margins from the oldest (morphotype 1) with a down-sloped anterior Quadralella spp. indet., and perhaps in Q. pardoneti. It appears that platform margin profile, to later forms (morphotypes 2, 3) witha elements with more ornate anterior margins arose independently from straight profile, followed by those (morphotype 4) with an elevated ancestors that had either a posterior pit (willistonensis > kathleenae), anterior node developed on one margin. Morphotype 3 is atypical in or one located more to the anterior (noah > mcrobertsi). A similar but its high crested blade-carina, and morphotype 5 is unique in having more pronounced differentiation of anterior nodes in Quadralella is a biconvex platform outline. Morphotypes 6 and 7 have increased believed to have also led to the more denticulate Primatella, some large platform curvature and arching, a longer free blade, and a more anterior elements of which have regressive low anterior nodes. pit. Several additional Quadralella species are identified by their Metapolygnathus dylani sp. nov. is interpreted to have evolved distinctive posterior platform shapes. Uncommon representative of from M. ex gr. communisti through reduction of the anterior platform the Q. oertlii (Kozur) group have triangular platforms that expand and the relative lengthening of the free blade. A continuation of progressively from the anterior to the broad posterior margin (Fig. this trend is thought to have led to M. parvus Kozur, which has a 9): two morphotypes are differentiated based on pit position and substantially reduced platform that is less than half of the total element anterior platform profile. A further species group that is subdivided length and a far anterior pit (Fig. 13). Three morphotypes of M. more formally into subspecies is Q. posteroexpansa sp. nov., which parvus are distinguished here: the alpha morphotype shows decreased has subparallel anterior platform margins that expand strongly in the platform length; the beta morphotype has reduced platform breadth; posterior one-quarter to one-half of the element (Fig. 10). Subspecies and the blade-like gamma morphotype has a completely reduced differ in the position of their pit, the position of the posterior platform platform. The latter was the basis for an earlier suggestion (Orchard, expansion, blade length, and anterior ornament. Two morphotypes of 1991a) that Norigondolella arose via such platformless morphotypes a further distinctive species, Q. deflecta sp. nov., are identified. These of Metapolygnathus, but this is no longer regarded as tenable because linguiform elements occur at two levels and show variation in posterior Norigondolella appears earlier. carina development (Fig. 10). Asymmetrical platforms are also a feature of the elongate and sinuous Q. sigmoidalis sp. nov., and of the shorter Parapetella (Figs. 14-16; 50-60) Q. roysi sp. nov. P1 elements assigned to Parapetella are quite variable but are united Kraussodontus (Figs. 11, 12; 42-45) in having distinctly raised parapets or buttress-like anterior platform margins that bear little or no additional ornament. Three subgroups of Kraussodontus was introduced for Late Carnian platform elements P1 elements are differentiated within the genus: those with relatively characterized by a relatively narrow, commonly elongate platform with short and broad platforms with variable posterior outlines (Fig. 14); a narrow pointed to broadly rounded posterior margin, and generally relatively elongate elements that have a medial platform constriction low and inornate, or weakly ornate anterior margins (Orchard, 2013). and a laterally expanded posterior platform (Fig. 15); and those with a Three species appear low in the BBR section, K. reversus (Mosher), relatively narrow, often tapered or pointed posterior platform (Fig. 16). K. peteri Orchard (Fig. 11), and K. ludingtonensis sp. nov. (Fig. 12). The present author has considered creating additional genera for these The P1 element of the first species has a generally low carina and lacks subgroups, but they are currently kept united in Parapetella. both a free blade and marked anterior geniculation points, whereas a The first subgroup includes the type species, Parapetella high anterior blade and marked geniculation points characterize K. prominens Orchard, which has high, very well differentiated anterior peteri, two morphotypes of which show progressive platform reduction margin parapets, and posterior platform outlines that vary from and lengthening of the free blade (Fig. 11). The same blade-platform subrectangular, to rounded, to incurved linguiform (Fig. 14). These characteristics are seen in the younger morphotypes of K. roberti variations are assigned to separate subspecies of P. prominens, all sp. nov., which differ in their submedial pit, more rounded posterior of which appear close to the CNB; only P. p. prominens subsp. nov. margins, and steep anterior platform margins. Longer incurved appears to range into younger strata. An equally short ranging element elements are assigned to K. rosiae sp. nov., and the strongly incurved linked to these taxa is referred to P. sp. nov. C, which has a strongly K. urbanae sp. nov. (Fig. 11). An exceptional specimen assigned to K. reduced platform. This suite of taxa stratigraphically succeeds elements praeangustus has the long carina, lower blade, and posterior pit like in which the parapets are less well differentiated but clearly raised with that seen in K. reversus, but it has geniculation points on the anterior respect to the posterior margins: these were called Metapolygnathus sp. margins. nov. Q by Orchard (2007c). The stratigraphic occurrences of several A subgroup of Kraussodontus, starting with K. ludingtonensis, allied species differ, so they are named individually: subrectangular has P1 elements with an elongate-oblong platform with a rounded to elements are referred to P. beattyi sp. nov., more tapered elements to subrounded posterior margin, and straight to slightly indented lateral P. broatchi sp. nov., and incurved linguiform elements to P. clareae margins (Fig. 12). K. ludingtonensis, with its spoon-like posterior sp. nov. Less common are abbreviated, posteriorly rounded elements platform, may have given rise to the more bowed K. aff. margaretae assigned to P. sp. nov. A, and elements with strongly reduced platforms through posterior carina growth and anterior shifting of the pit. Younger assigned to P. sp. nov. B (Fig. 14). K. margaretae is broader and shorter, and shows further anterior pit The second subgroup of Parapetella species appear later than migration. Both K. vancouverense sp. nov. and K. wendae sp. nov. the first, and all have medial to anterior pits (Fig. 15). Two separate 11

Figure 8. Proposed phyletic linkages in large subrectangular elements of Quadralella with low, variable nodes, and a pit that migrates anteriorly in some (on right) through time. See Figure 7 for legend. 12

Figure 9. Proposed phyletic linkages in elements of Quadralella with a large, broad platform and few nodes; pit migrates anteriorly, and free blade may become longer (on right) through time. See Figure 7 for legend. 13

Figure 10. Proposed phyletic linkages in medium to large size species of Quadralella with an asymmetrical posterior platform; platform size diminishes and pit migrates anteriorly through time. See Figure 7 for legend. 14

Figure 11. Proposed phyletic linkages in species of Kraussodontus with a rounded to pointed, straight to curved platform; free blade lengthens/ platform shortens (on left), and pit migrates anteriorly through time. See Figure 7 for legend. 15

Figure 12. Proposed phyletic linkages in species of Kraussodontus with a relatively flat and elongate platform that may have a submedial constriction (on right); pit migrates anteriorly through time. See Figure 7 for legend. 16

Figure 13. Proposed phyletic linkages in species of Metapolygnathus with a generally inornate platform bearing a submedial to anterior pit; anterior pit migration, and strong platform reduction (on right) may occur through time. See Figure 7 for legend. 17

Figure 14. Proposed phyletic linkages in species of Parapetella with a variable platform shape but showing increased differentiation of the anterior parapets through time. See Figure 7 for legend. 18

Figure 15. Proposed phyletic linkages in species of Parapetella with a medial platform constriction: some show a posterior expansion (on left); others marked platform reduction (on right) through time. See Figure 7 for legend. morphological trends are recognized in this group: one involving rise to weakly posteriorly expanded P. irwini sp. nov., the strongly progressive platform reduction; the other parapet growth and posterior expanded P. posterolata sp. nov. Much later, the similarly shaped P. expansion. The oldest representative is P. elegantula sp. nov., sp. nov. G with its more differentiated parapets and a more anterior pit characterized by a subparallel-sided P1 element with a slight medial appeared. narrowing and bulbous posterior platform (Fig. 15). This species The third subgroup of Parapetella includes P1 elements with is hypothesized to have given rise through platform reduction to P. tapered, narrowly rounded to pointed posterior platforms, and lanei sp. nov., in which a stronger mid-platform constriction separates rounded to pointed anterior platform margin parapets (Fig. 16). The the raised anterior margins from a low and expanded posterior part. oldest representative is P. aff. riteri, which has an inner parapet but a Through further narrowing of the platform of P. lanei, both the posterior nodose outer margin, as well as a relatively posterior pit. Two partly and particularly the anterior platform becomes vestigial or absent, as in co-occurring morphotypes of the narrowly tapered P. riteri sp. nov. the diminutive P. pumilio sp. nov. and P. aff. pumilio. In the slightly have differing shaped parapets: rounded in the alpha morphotype and younger, equally diminutive P. johnpauli sp. nov., the hourglass relatively angular in the beta morphotype (Fig. 16). Younger strata platform shape of P. lanei is retained, but the blade lengthens and the contain the diminutive P. willifordi sp. nov., which has a pointed pit lies at the anterior end of the platform. In P. sp. nov. D, only a platform like that of P. riteri, but is reduced in size, has a longer blade, small, subtriangular posterior platform is retained (Fig. 15). Concurrent and an anterior pit. Sharply terminated parapets and an abbreviated with strong platform reduction in one branch of this group, the ancestral platform also characterize the older P. rubae sp. nov. Parapetella elegantula is hypothesized to have broadened and given A separate trend in this third subgroup of Parapetella is towards 19

Figure 16. Proposed phyletic linkages in species of Parapetella that have a narrow to pointed posterior platform, and show marked reduction in either platform width (on right) or length (on left) through time. See Figure 7 for legend. 20 elongation and then increased narrowing of the posterior platform then develop more on the lateral margins in front of a constriction that starting with P. columbiense sp. nov., and leading to P. cordillerense migrates from a posterior position, as in C. eozoae Orchard to a more sp. nov. (Fig. 16). Overall platform reduction in the contemporaneous central position, as in C. zoae (Orchard). A possible ancestor (from Haida and possible derivative P. hillarae sp. nov. follows a trend of Gwaii) is shown as C. aff. eozoae in Figure 17: this form may succeed platform shortening and subsequent narrowing to produce the small, C.? lindae (Orchard). Concurrent with the transition from C. eozoae to subrectangular posterior platforms and anterior parapets manifest in C. zoae, some elements exhibit both rounded nodes and sharp denticles variants or morphotypes of the diminutive P. destinae sp. nov. This on their platform margins: these are differentiated as C. anitae sp. nov. subgroup of Parapetella shows the same pattern of platform reduction As originally conceived, Carnepigondolella was wholly Late observed in some Acuminatella species, which differ in having sharp Carnian and culminated in relatively diminutive elements now referred anterior nodes rather than parapets. to C. spenceri, C. aff. spatulata, and possibly C. echinatus. An identical trend toward diminutive faunas is repeated in both Metapolygnathus Carnepigondolella (Figs. 17, 18; 35-38) and Parapetella at the CNB (see above). The two events may have Kozur (2003) introduced Carnepigondolella for Late Carnian been confused in the past, but at BBR they are clearly separated by ornate taxa, with C. zoae (Orchard) as the type species. The present author strata that document the ascent of Acuminatella and Primatella in the (Orchard, 1991a, pp. 176, 182) suggested that constituent species of this absence of Carnepigondolella. Assignment of several other younger lineage (then called Metapolygnathus n. sp. E, M. sp. F, M. sp. G, and species to Carnepigondolella (e.g., Mazza et al., 2012b) obscures what M. echinatus Hayashi) showed progressive heightening and sharpening is regarded as a potentially useful datum in Late Carnian conodont of the anterior platform nodes and, finally, reduction of the platform history. length. This proposed lineage may have been an oversimplification because species with small and sharp denticles are present throughout Acuminatella (Figs. 19, 20; 32-34) the lower part of the BBR section, prior to the appearance of those with The type species of this genus, Acuminatella acuminata Orchard, rounded nodes and mixed ornament. The present section does not reveal has a posterior platform that is regularly tapered, a relatively straight the early evolution of these forms, which remains unclear. However, carina that extends to its pointed posterior end, and sharp marginal the youngest representatives of Carnepigondolella do indeed show anterior denticles. It is preceded by the shorter A. sagittale sp. nov., progressive platform reduction, and may include the species echinatus which has less differentiated, apically rounded nodes on the anterior (see Carter and Orchard, 2013). It may be useful in the future to restrict margins of the P1 platform elements (Fig. 19). Similarly shaped platform Carnepigondolella to taxa with well-developed rounded nodes, like C. elements that entirely lack nodes (or parapets, as in Parapetella) are zoae and its suggested precursor, and establish a new genus for sharply assigned to Kraussodontus, species of which may be ancestral to the ornate forms like C. samueli. The opposite approach has been taken by sagittate Acuminatella species. Mazza et al. (2012b), who expanded the genus to include some taxa Acuminatella sagittale is the oldest species unequivocally with relatively subdued nodes (referred to Quadralella in this work). assigned to the genus and provides a starting point for a morphological The stratigraphically oldest representatives of Carnepigondolella trend that leads to A. constricta sp. nov., which has similar anterior at BBR are assigned to C. samueli (Orchard) and C. gibsoni sp. nov., nodes but a posterior platform that is reduced to a narrow flange on which have similar sub-rectangular platforms but differ in that the latter each side of the well-developed carina. Anteriorly sharp nodes first species has no, or a very subdued, posterior platform ornament (Figs. appear in the small species A. binodosa sp. nov., which probably 17, 18). Among the more ornate group, C. samueli and the successive includes juvenile specimens of partly contemporaneous Acuminatella C. postsamueli occur as both symmetrical and posteriorly asymmetrical species, such as A. acuminata and A. denticulata sp. nov. Similarly, morphotypes. The reduction in platform length and concomitant blade A. binodosa may include early growth stages of Primatella species, lengthening culminates in relatively small and short-ranging elements and later, juvenile Epigondolella. This small species is a homeomorph assigned to C. aff. spatulata (Hayashi), in which platform and blade are of Late Norian E. bidentata, the recognition of which poses similar about equal in length (Fig. 17). problems. Acuminatella denticulata differs from its presumed ancestor, Contemporaneous Carnian taxa that lack distinctive posterior A. constricta, in bearing sharp anterior denticles, repeating the trend ornament are referred to Carnepigondolella gibsoni, and later to two seen in the transition from A. sagittale to A. acuminata. The Norian A. morphotypes of C. pseudodiebeli (Kozur) (Fig. 18). This succession of curvata sp. nov. is thought to be a derivative of A. acuminata, differing species show relative broadening and shortening of the platform. The in its long axis curvature and broader platform. beta morphotype of C. pseudodiebeli has a platform shape reminiscent Additional species assigned to Acuminatella share the relatively of the much younger Epigondolella, but both the size/height of the long and slender platforms, increasingly sharp anterior denticles, anterior denticles, pit position, and lower surface profile generally unornamented posterior platform, and long carina of the type species, differ. These and similar elements may have formed the basis for but they also have the abruptly narrowed posterior platform (Fig. 20) suggesting a record of Epigondolella low in the Pizzo Mondello (PM) and may have had a different origin. The oldest of these, A. sinuosa section in Sicily (Mazza et al., 2012b), much earlier than appearance of sp. nov., has a narrow incurved posterior platform, a morphology that Epigondolella sensu stricto in British Columbia. persists through much of the higher BBR section. Derivative species Overlapping with the upper range of Carnepigondolella show increased reduction in the length of the anterior platform and pseudodiebeli are several new, posteriorly inornate taxa that have a relative lengthening of the posterior platform: this reaches an extreme broadened or rounded posterior platform, as well as longer blades: development in the beta morphotype of A. angusta Orchard and the C. milanae sp. nov., C. spenceri sp. nov., and C. aff. spenceri (Fig. bidentate A. longicarinata sp. nov., a homeomorph of the Rhaetian 18). A marked medial narrowing followed by posterior expansion of Epigondolella mosheri (Kozur and Mostler). the platform is a diagnostic platform shape for the contemporaneous The origin of these elongate and posteriorly reduced Acuminatella C. medioconstricta Orchard. The diminutive C. echinatus (Hayashi), species may lie in the slightly older A. sagittale sp. nov. through abrupt although not recovered from BBR (but see Carter and Orchard, 2013), narrowing of the posterior platform, as proposed for A. constricta. may also be a member of this diverse group, in this case derived through However, the acquisition of small but sharply terminated anterior the total reduction of the anterior platform to produce a form with an denticles seen in A. sinuosa occurred earlier than in the A. acuminata anterior rather than medial pit. Rare specimens of Carnepigondolella? subgroup. Preceding A. sinuosa stratigraphically are uncommon sp. nov. A may represent an earlier product of platform reduction species questionably included in Acuminatella, namely A.? prima in this stock: its denticulation is identical with contemporaneous sp. nov. and A.? extensa sp. nov. (Fig. 20). These have very similar Carnepigondolella species but it is unique in having a pointed platform, but more numerous anterior marginal nodes compared with typical similar to that of the younger Acuminatella. This author does not regard Acuminatella, but they differ in having a broader posterior platform, and C.? sp. nov. A as an evolutionary step to Acuminatella because the first a carina that terminates in an enlarged cusp surrounded by a posterior representatives of the latter genus lack a posterior carina, medial pit, platform brim. They share the latter features with the even older and and sharp denticles, all features developed later. possibly ancestral Quadralella postlobata, in which differentiation All the aforementioned taxa currently assigned to Carnepigondolella of the anterior platform nodes is variable. This alternate derivation, are united in their small but sharply terminated platform denticles. In shown in Figure 20, would involve progressive anterior shifting of the contrast, the genus, as exemplified by the type species C. zoae, differs platform constriction, concomitant elongation and attenuation of the in its well defined but low circular nodes when viewed from above. posterior platform, and further development of a posterior carina, as Rounded nodes are first introduced at the anterior platform margin and well as increased development of the anterior denticles. 21

Figure 17. Proposed phyletic linkages in species of Carnepigondolella, including ornate elements with rounded nodes (on right), and quadrate forms with sharp denticles on all margins and diminishing platform length (on left) through time. Illustrations marked “HG” are from Haida Gwaii. See Figure 7 for legend. 22

Figure 18. Proposed phyletic linkages in species of Carnepigondolella with well-developed denticles confined to the anterior platforms; platform length also diminishes through time. Illustration marked “HG” is from Haida Gwaii. See Figure 7 for legend. 23

Figure 19. Proposed phyletic linkages in species of Acuminatella with pointed platforms and increased anterior denticle development through time, and some with posterior platform reduction (on right). See Figure 7 for legend. 24

Figure 20. Proposed phyletic linkages in species of Acuminatella with a medial constriction that is emphasized as the posterior platform narrows and lengthens through time; anterior nodes also become sharper. See Figure 7 for legend. 25 Primatella (Figs. 21-23; 61-74) platforms (Fig. 25). In all cases, the oldest examples have apically rounded and less differentiated anterior nodes. Elongate elements of Virtually all species of Primatella were regarded as variations P. oblonga sp. nov. and broader elements of P. rotunda sp. nov. are within the Epigondolella primitia population of Orchard (1983). Since differentiated as alpha and beta morphotypes based on their anterior then, there has been much uncertainty about the identity of similar node formation, and in the case of P. rotunda, a pit position that migrates elements occurring in North America and in Tethyan regions, and this to the anterior. The posterior platform of P. asymmetrica Orchard has a has led to a much narrower definition of species. Those introduced here convex outer margin and a more or less straight inner margin. Typical are similarly narrow in scope. This approach is aimed at both improving specimens are preceded by narrower forms with much lower anterior correlations between, and potentially differentiating faunas from, either denticles, here called P. aff. asymmetrica. The similarly shaped but side of the Panthallassa Ocean. diminutive P. sp. nov. A is rare in the upper Kerri Zone near the top of The holotype of Primatella primitia has a subrectangular platform the BBR section. and three closely spaced carinal nodes in front of a broad platform brim (Fig. 21). Such forms are relatively rare at BBR, but they occur through Epigondolella (Figs. 22-25; 61-74) almost the entire range of the genus. More common are similarly shaped The type species of Epigondolella is E. abneptis (Huckriede, platforms with two widely spaced denticles on the posterior platform, 1958). The holotype, now lost, is known to have come from Middle which are now assigned to P. conservativa Orchard, and others with Norian (Alaunian) strata in Austria (Kozur, 2003; Moix et al., 2007, several posterior carinal nodes that almost reach the posterior margin, p. 291), but it has never been well described. In the past, the species here referred to P. elongata sp. nov. Later, broader rectangular elements has been uncritically identified in Upper Carnian and, most commonly, with an indentation in the posterior margin appear and these are in the Lower Norian, but with the introduction of the more precisely described here as P. bifida sp. nov. defined E. quadrata Orchard, and of Carnepigondolella, use of E. Apart from the relatively long, rectangular taxa noted above, abneptis has declined and modern reports of the species lack clear Primatella species may be characterized by a variety of other platform meaning. shapes: short and broad quadrate; short and tapered; rounded or Epigondolella quadrata is regarded by the present author as asymmetrical posterior outlines; and platforms with a distinctive medial the first species of the genus, which is consequently regarded asno constriction and expanded posterior platform. It is believed that all these older than Lower Norian. Morphological variation in the faunas was elements arose from ornate Quadralella species, like Q. postlobata, in presented as the E. abneptis subsp. A population by Orchard (1983), which anterior nodes became increasingly differentiated and sharply and the multiple species of the genus now identified at BBR were terminated. The earliest examples of Primatella have anterior nodes/ formerly combined therein. Notably, all these Epigondolella taxa appear denticles of intermediate development, as in P. rectangulare sp. nov. simultaneously at BBR, where they abruptly replace the spectrum of and P. mclearni sp. nov. Primatella species (Table 8). This contrasts with the situation at Pizzo Many early growth stages of Primatella are medially restricted, Mondello (PM), Sicily where several apparently older species are but those that retain such morphology into later growth stages are assigned to Epigondolella (e.g., Mazza et al., 2012a, b). Such records separated in this work. Elements grouped in the oldest of these, P. challenge the hypothesis presented here that Primatella was the direct mclearni, have a variable carina and probably represent precursors to ancestor of Epigondolella. As illustrated in this work, similar platform both P. orchardi (Kozur) with its centrally located posterior cusp, and shapes are present in both Primatella and Epigondolella, which can be the broadly contemporaneous P. stanleyi sp. nov., in which additional distinguished by the much higher and sharper anterior denticles of the posterior carinal nodes occur (Fig. 22). Two further derivatives latter. Hence, E. quadrata resembles P. orchardi (Fig. 22), elements show either a reduction in posterior platform width, as in Primatella compared with E. miettoi and E. vialovi resemble E. subquadrata (Fig. posteroglobosa sp. nov., or broad expansion of the posterior margin, 23), E. aff. uniformis resembles P. permica (Fig. 23), E. sp. nov. A as in P. vanlierae sp. nov. These two species have a broadly similar shares attributes of the P. ovale (Fig, 24), and E. aff. stefanionensis platform shape as Parapetella lanei and P. posterolata, but differ in has the shape reminiscent of P. rotunda (Fig. 25). It is not clear at having anterior denticles. A further variation on the medially constricted present whether Tethyan elements assigned to Epigondolella prior to forms is Primatella circulare sp. nov., in which the posterior platform is the appearance of E. quadrata sensu stricto are actually representatives semicircular. of Primatella, or even the older Carnepigondolella. Thorough revision Primatella species with a relatively short rectangular platform of Epigondolella is best achieved through study of rich Norian faunas start with P. rectangulare. It shows poorly differentiated but sharply that post-date the CNB interval. terminated anterior nodes in contrast with the round-topped nodes of similarly shaped elements of its possible forbearer Q. postlobata (Fig. CONODONT ZONATION 23). Similarly shaped elements with more differentiated and sharp The first biostratigraphic summary of the BBR section (Orchard et anterior denticles and several posterior carinal nodes are included in P. al., 2001b, c) incorporated the prevailing conodont taxonomic concepts mersinensis (Kozur and Moix), whereas those with a central terminal presented by Orchard (1983, 1991a, b). More detailed sampling and cusp are referred to P. subquadrata sp. nov. Later, large specimens taxonomic revisions in the next decade enabled subdivision into a of the posteriorly expanded P. triangulare sp. nov. appear. A further number of informal conodont zones (Orchard, 2007c; 2010b, 2013) variation on these subquadrate elements is P. permica (Hayashi), which based partly on new and undefined taxa. With the definition of these taxa shows the first development of posterior ornament in Primatella, a in this paper, the zones are formally defined (Fig. 4): their occurrences feature that becomes common in the younger derivative Epigondolella. in the outcrop are shown in Figure 26. In this account, conodont zones Relatively abbreviated, more or less tapered Primatella species, are distinguished from ammonoid zones by the use of italicized species generally with posterior platform brims developed posterior of discrete names for the former, e.g., primitia Zone versus Kerri Zone. cusps, are uncommon and rather variable (Fig. 24). Squat elements that The utility of the original primitia Zone, based on the range have a length-to-breadth ratio of between 1:1 and 1:1.5 are combined of the “Epigondolella primitia population” (later assigned to as the P. pseudoechinata (Kozur) group, whereas those with narrower, Metapolygnathus, and now Primatella), is reflected in the retention of more pointed platforms are referred to P. ovale sp. nov. Both may be an amended Primatella primitia Zone. This zone is well known, readily related to the older Quadralella stephanae (Orchard). A single element identifiable, and clearly encompasses the traditional ammonoid-based from PHE closely resembles the holotype of P. pseudoechinata in its CNB. Subdivision of the primitia Zone into “Lower” and ”Upper” relatively rounded platform, but elements from various stratigraphic parts is abandoned, and new subzones are introduced, including levels at BBR are more tapered or angular in outline; the youngest the re-cast parvus Subzone, and a lowermost part formerly called example is unique in having an inner parapet. Collectively, these the “Metapolygnathus n. sp. Q Zone.” Similarly, the samueli Zone, elements may be amenable to further speciation but they are too few originally introduced as a subzone of an ill-defined Metapolygnathus in the present collections to do so. The oldest of these abbreviated nodosus Zone (Orchard, 1991a), is elevated to a range zone that is elements, referred to P. aff. ovale, has less sharp anterior platform divisible into at least three subzones. denticles, a shorter posterior platform, and a more posterior pit than The new zonation is listed in descending stratigraphic order below, younger elements assigned to P. ovale. Theoretically, the addition of and then described in ascending order: low marginal nodes on the posterior margin of P. ovale would yield P. rhomboidale sp. nov., a probable Norian derivative. Epigondolella quadrata Zone A final subgroup of Primatella includes elements with relatively Primatella primitia Zone broad, terminally rounded, symmetrical to asymmetrical posterior Primatella asymmetrica-Norigondolella sp. Subzone 26

Figure 21. Proposed phyletic linkages in species of Primatella with an elongate-rectangular platform with strong anterior denticles, and variable posterior carinas. See Figure 7 for legend. Metapolygnathus parvus Subzone co-occurring, these taxa are assigned to the new zoae Subzone of the Acuminatella acuminata-Parapetella prominens Subzone samueli Zone. Several other species of Carnepigondolella have a more Acuminatella angusta-Metapolygnathus dylani Subzone restricted range within the samueli Zone, and enable subdivision of the Acuminatella sagittale-Parapetella beattyi Subzone zone into a total of four subzones: these are based on the successive Carnepigondolella samueli Zone appearances of C. eozoae, C. zoae, C. medioconstricta, and C. spenceri. Carnepigondolella spenceri Subzone In addition to Carnepigondolella species, more common but longer Carnepigondolella medioconstricta Subzone ranging elements of the samueli Zone are species of Kraussodontus Carnepigondolella zoae Subzone and Quadralella. Elements of Q. carpathica invariably dominate Carnepigondolella eozoae-K. ludingtonensis Subzone collections in this zone, and are often accompanied by Q. lobata and Q. posteroexpansa alpha morphotype, and later by K. ludingtonensis Carnepigondolella samueli Zone (beds P-4). and Q. angulata; K. peteri alpha morphotype, K. aff. margaretae, K. This zone is defined by the range of Carnepigondolella samueli reversus, Metapolygnathus ex gr. communisti, and Q. ex gr. oertlii occur (Appendix, Table 3). At BBR, the entire exposed Ludington Formation rarely. The lowest parts of the samueli Zone lacks subzonal indicators is assigned to this zone, although neither the base of the zone nor that of and is undifferentiated (Fig. 3). The samueli Zone is a minimum of the formation are seen. In Haida Gwaii (Fig. 1, inset), Orchard (1991a, b) about 50 m thick at BBR. introduced a samueli Subzone and an older zoae Subzone in the middle to upper parts of a broadly defined nodosus Zone. However, recent Carnepigondolella eozoae-Kraussodontus ludingtonensis Subzone taxonomic re-assessment of the type of “Metapolygnathus” nodosus (beds K-A). does not favour its use in Upper Carnian zonation (Moix et al., 2007, The lowest 15 m of the exposed Ludington Formation contains p. 291). The two species of Carnepigondolella were thought to succeed Carnepigondolella samueli and C. gibsoni and these strata are therefore one another in Haida Gwaii, although their holotypes came from a single assigned to undifferentiated samueli Zone. The two nominal species, collection. At BBR, C. samueli appears earlier than C. zoae, and when C. eozoae and Kraussodontus ludingtonensis, appear concurrently in 27

Figure 22. Proposed phyletic linkages in species of Primatella with a marked medial constriction and a variably expanded and carinate posterior platform. Epigondolella quadrata appears high in the section and has a similar platform shape but, as with all Epigondolella species, has much larger denticles. See Figure 7 for legend. bed K, thereby defining a base for this subzone, which is a minimum Subzone. Morphotype 1 of Metapolygnathus ex gr. communisti is also of 22 m thick at BBR (Appendix, Table 3). The eozoae-ludingtonensis only known from this subzone, but it is very rare. All other associated Subzone corresponds to part of Fauna 1 of Orchard (2010b, fig. 4). taxa range into the higher samueli Zone, including Q. angulata, Q. Above the base of this subzone, the index species are joined lobata, and abundant Q. carpathica. by Carnepigondolella pseudodiebeli and all three ornate taxa range into, or beyond, the supra-adjacent zoae Subzone. Both Quadralella Carnepigondolella zoae Subzone (beds 0d-1). posteroexpansa alpha morphotype and the uncommon beta morphotype The appearance of Carnepigondolella zoae defines the base of C. pseudodiebeli disappear at, or close to, the base of the zoae Subzone of a medial subzone of the samueli Zone. This subzone, formerly and therefore may be useful proxies for the eozoae-ludingtonensis included in Fauna 1 of Orchard (2010b), is ~10 m thick at BBR. The 28

Figure 23. Proposed phyletic linkages in species of Primatella with a relatively short subquadrate platform without marked constriction; one develops posterior ornament, another strongly expands laterally. Similarly shaped Epigondolella species appear high in the section. See Figure 7 for legend. 29

Figure 24. Proposed phyletic linkages in species of Primatella with a short ovoid platform that tapers to the posterior. A similarly shaped Epigondolella species appears high in the section. See Figure 7 for legend. primary index may be accompanied by C. anitae and rare examples conodont history with the appearance of the index species plus C. of Carnepigondolella? sp. nov. A, Kraussodontus aff. margaretae, and postsamueli, C. milanae, C. aff. spenceri, and Quadralella postlobata. Morphotype 2 of Metapolygnathus ex gr. communisti. Several species Each of these species are characterized by relatively long free blades disappear just above the base of the zoae Subzone, including C. gibsoni, in contrast to their proposed forebears, and the Quadralella species the beta morphotypes of both C. pseudodiebeli and C. samueli, and also have relatively raised and more ornate (noded) anterior platform later Quadralella lobata. Those Quadralella species known from the margins unlike the flattened lateral profiles of older representatives. eozoae-ludingtonensis Subzone continue to dominate faunas in this Kraussodontus peteri alpha morphotype is more common and also subzone (Appendix, Table 3). shows the beginning of blade lengthening at the expense of platform Carnepigondolella medioconstricta Subzone (beds 1c-2a). reduction. In addition, the appearance of Quadralella noah marks the beginning of anterior pit migration in the genus. Finally, the first, The appearance of Carnepigondolella medioconstricta defines the albeit questionable, Acuminatella species, A.? prima, appeared in this base of this subzone, which was formerly called the C. n. sp. N Zone subzone. (Orchard, 2007c), and later assigned to Fauna 2 of Orchard (2010b, fig. In spite of these morphological innovations, Quadralella is still 4). The medioconstricta Subzone, which is ~3 m thick at BBR, marks represented by abundant Q. carpathica and Q. angulata, common a significant diversification and evolutionary event in Late Carnian Q. ludingtonensis, and rare Q. oertlii. However, the newcomer, Q. 30

Figure 25. Proposed phyletic linkages in species of Primatella with an elongate, posteriorly rounded or asymmetric platform; anterior ornament becomes increasingly differentiated through time. A similarly shaped Epigondolella species appears high in the section. See Figure 7 for legend. postlobata, becomes a major component of the faunas in this subzone. rare examples of early Primatella species assigned to ?P. mclearni and A single specimen of Misikella longidentata Kozur and Mock was P. aff. asymmetrica, plus distinctive elements assigned to Quadralella also recovered from this subzone, which saw the last occurrence of C. mcrobertsi, and the alpha morphotype of Q. deflecta. The first three anitae, C. zoae, and K. reversus (Appendix, Tables 3, 4). taxa exhibit increased differentiation of anterior platform nodes seen Carnepigondolella spenceri Subzone (beds 3-4). also in some contemporary Q. postlobata, the proposed root stock. As is the case throughout the samueli Zone, Q. carpathica and related species A thin, ~1 m interval at the top of the samueli Zone is distinguished remain abundant, but to these are added early examples of Q. tuvalica. here as an uppermost subzone defined by the range ofCarnepigondolella Rare morphotypes 2 and 3 of Metapolygnathus ex gr. communisti spenceri. The interval was originally characterized (erroneously) as the also occur in this subzone. By the end of the spenceri Subzone, and Carnepigondolella pseudoechinata Zone by Orchard (2007c), and later of the samueli Zone, Carnepigondolella (sensu herein) disappeared as Fauna 3 by Orchard (2010b). Additional guides for this uppermost (Appendix, Tables 3, 4). subzone are C. aff. spatulata and Quadralella karenae; C. echinatus sensu stricto may also belong to this fauna (see Carter and Orchard, Primatella primitia Zone (beds 5-31). 2013). All these species are characterized by platform diminution, An Epigondolella primitia Zone was not differentiated when Sweet continuing the trend begun in the medioconstricta Subzone: among et al. (1971) first summarized Triassic conodont zonation, although the those elements assigned to C. aff. spatulata, this is apparent even within unnamed species was included in their Carnian Zone 19. Later, after the subzone (Fig. 17). Mosher (1973) named the species, it was recognized that the species Making a first appearance near the top of the samueli Zone are straddled the CNB, and a zone of that name was warranted. Orchard (1983, 1991a, b) subdivided the primitia Zone into lower and upper 31

Figure 26. Photograph of the lower Black Bear Ridge section showing the assignment of strata to conodont zones (above) and subzones (below). The CNB interval delineated by the white rectangle is shown in Figure 27. parts based on the appearance of Norigondolella navicula midway those of contemporaneous inornate Kraussodontus and Quadralella through the zone. The latter species, or allied ones, are abundant in some species. Some associated Parapetella species that appear in the Norian beds of the Pardonet Formation (Fig. 3), but rare specimens subzone have very short (P. sp. nov. A and P. sp. nov. B), or asymmetric appear earlier. The origin of Norigondolella remains uncertain, but its platforms (P. clareae). The rare occurrence of P. aff. riteri foreshadows bloom in the Lower Norian is probably environmentally controlled. further diversification of the genus in the overlying subzone. Co- Orchard (2013) more recently differentiated lower and upper parts of occurring Kraussodontus peteri beta morphotype, and later K. roberti the primitia Zone based on position relative to the intervening parvus alpha morphotype, also have a reduced platform compared with the Subzone, but this too is superseded by the subzonal divisions of an older Kraussodontus. expanded primitia Zone. The sagittale-beattyi Subzone also marks the first occurrence of The base of the primitia Zone is here defined as the appearance the ornate taxa Acuminatella sinuosa, A.? extensa, Primatella mclearni, of Acuminatella and common Primatella species in the absence P. oblonga alpha morphotype, and P. rectangulare. Although less of Carnepigondolella. Species of both the former genera become common than contemporaneous inornate elements, these and related increasingly common in younger faunas and then become dominant elements become increasingly common and diversified up-section. in the Norian (Fig. 6). A single ?P. mclearni is known immediately Their appearance in this subzone provides a useful marker for the base prior to the primitia Zone, but the associates are quite different. Many of the primitia Zone. Primatella species have a long range through the primitia Zone and Quadralella is represented in this subzone by the last Q. angulata, extend up to the appearance of Epigondolella quadrata, which marks Q. carpathica, and Q. ludingtonensis. Both Q. postlobata and Q. its top. The uppermost subzone of the primitia Zone is quite different tuvalica are common, whereas Q. deflecta alpha morphotype, Q. from those in its lower part by lacking many of the inornate taxa. The stephanae, and Q. roysi are rare. Numerous small elements assigned to entire primitia Zone is ~34 m thick at BBR. Q. sp. indet. appearing at this level are probably early growth stages of some of these species. Metapolygnathus ex gr. communisti morphotype Acuminatella sagittale-Parapetella beattyi Subzone (beds 5-11c). 4 also occurs rarely (Appendix, Table 4). Formerly referred to as the “Metapolygnathus n. sp. Q Zone” by Acuminatella angusta-Metapolygnathus dylani Subzone (beds 12- Orchard (2007c, 2013), and to Fauna 4 by Orchard (2010b), this ~5 16). m interval heralds a major change in the conodont fauna following the disappearance of Carnepigondolella, the abundant appearance This subzone is equivalent to the larger part of the Metapolygnathus of Parapetella, and the increasingly common occurrence of both n. sp. P - n. sp. Y Zone of Orchard (2007c), and the “Lower primitia Acuminatella and Primatella. The nominal indices represent the first of Zone” of Orchard (2013); it also corresponds to Fauna 5 of Orchard their respective genera, and the faunal turnover that introduces them is (2010b). About 8 m in thickness, these strata yield an abundant and judged to be second only to that within the parvus Subzone (see below). diverse conodont fauna dominated by species of Kraussodontus, The appearance of Acuminatella sagittale, which is characterized Parapetella, and Quadralella, and fewer Metapolygnathus. Many of by a long carina and a tapered and pointed platform, coincides with that these can be grouped into clades that show progressive evolutionary of Parapetella beattyi, which has distinctly raised and unornamented morphogenesis that can be traced through the parvus Subzone, where anterior platform margins. Platform profiles of the latter contrast with many disappear (e.g., Figs. 13-16). About 20% of elements are assigned 32 to ornate species of Acuminatella and Primatella, many of which are Orchard (2013). Fifty-eight taxa range into this subzone from underlying long-ranging. Unique occurrences of Norigondolella sp. and Misikella strata, a further 19 appear, and 16 have their last occurrence within the sp. nov. A also occur (Appendix Tables 4, 5). subzone (Fig. 6, Appendix, Tables 4-7). This represents both a period The species chosen as nominal taxa for the angusta-dylani of evolutionary innovation, and the initial die-off of long-ranging Subzone are distinctive, but are not the most common. Nevertheless, Carnian taxa that reaches its peak midway through the overlying parvus Acuminatella angusta is abundant and the increase in length and Subzone. reduction in breadth of the posterior platform in the alpha morphotype Acuminatella acuminata, which appears at the base of the reaches an extreme in the shorter ranging beta morphotype. The less subzone, and Parapetella prominens subspp., which appear a little common Metapolygnathus dylani is believed to have developed higher, are chosen as nominal indices: the occurrence of either taxon through platform shortening from Metapolygnathus ancestors with prior to appearance of Metapolygnathus parvus identifies the subzone. longer platforms, none of which are common in the Canadian faunas; Also significant are the first appearances of Primatella oblonga beta morphotype 4 of M. ex gr. communisti occurs contemporaneously near morphotype, P. permica, P. triangulare, Parapetella irwini, and the origin of M. dylani. P. posterolata. Compared with their predecessors, species of the In addition to Acuminatella angusta, the species A. sagittalis and acuminata-prominens Subzone have more strongly differentiated its derivative A. constricta range through the subzone, and these are anterior nodes, as in Acuminatella (acuminata, denticulata); or accompanied by the diminutive A. binodosa and A. longicarinata. Both parapets, as in Parapetella (irwini, posterolata, prominens); or greater the latter species could include early growth stages of other Acuminatella expansion of the posterior platform (triangulare); or posterior platform species, but it cannot be ruled out that some represent independent end- ornamentation (permica). Other notable first appearances are of members resulting from platform reduction. All of these taxa other Parapetella destinae alpha morphotype, P. pumilio, and P. sp. nov. C, than the beta morphotype of A. angusta range throughout the subzone all of which show the diminutive trend that becomes common in the and beyond. overlying parvus Subzone. Twelve Primatella species make their first appearance in the As noted, most of the species that appeared in the underlying angusta-dylani Subzone and most range through the remainder of sagittale-beattyi Subzone range into the present subzone. Quadralella the primitia Zone. These species can be divided into elongate forms praecommunisti subspp. remain very important and are often dominant typified byP. conservativa, P. elongata, and P. primitia; subrectangular in the collections from the acuminata-prominens Subzone. Other related forms such as P. mersinensis and P. subquadrata; and medially but less common elements become additions to the fauna, namely the constricted forms like P. orchardi, P. posteroglobosa, P. stanleyi, and beta morphotypes of both Q. deflecta and Quadralella pardoneti, and P. vanlierae. These are accompanied by less common examples of the Q. posteroexpansa subsp. B. Elements of Kraussodontus praeangustus more primitive and probable forebears represented by the quadrate P. and Quadralella? sp. nov. A occur rarely, as does Norigondolella sp. rectangulare and constricted P. mclearni. Less common species are the Two specimens of the latter occur (Table 6), but neither are clearly more rounded P. oblonga and P. rotunda, which both exhibit increasing examples of N. navicula. differentiation of the anterior nodes towards the top of the subzone, and the relatively squat P. aff. ovale and P. ex gr. pseudoechinata. Those Metapolygnathus parvus Subzone (beds 18c-21f). Primatella species that make a first appearance in the subzone occur in This interval is defined as the range of the nominal species and a staggered and sporadic fashion whereas later, when the species are is synonymous with the Metapolygnathus echinatus Zone of Orchard more common, they often co-occur. The speciation presented here goes (2007c), and Fauna 7 of Orchard (2010b). The interval is ~3 m thick beyond the “splitting” already presented in the literature and is designed at BBR and is readily recognized by the presence of the nominal to maximize their stratigraphic and biogeographic potential. However, index species and later by additional diminutive taxa of Parapetella none of the common species appear to have narrow stratigraphic ranges; (Appendix, Tables 6, 7). The appearance of these forms coincides with whether any are limited geographically remains to be determined. the disappearance of most other inornate elements (Fig. 6), and so the Ten Parapetella taxa appear in the angusta-dylani Subzone. Those short-ranging diminutive elements become a significant part of the that appear in the underlying subzone (P. beattyi, P. clareae) range fauna alongside the ornate Acuminatella and Primatella species that upward and are joined by P. broatchi. A second group consists of P. continue on through the boundary interval. The diminutive elements elegantula and more common P. lanei, which both display elongate, disappear at the top of the parvus Subzone, at which level the former medially constricted platforms. A third group, characterized by tapered genera become overwhelmingly dominant. Conodont collections platforms of variable length, consists of common P. columbiense, P. from this subzone provide a distinctive marker in the CNB interval of cordillerense, P. hillarae, and P. riteri alpha and beta morphotypes, and northeast British Columbia. rare P. aff. destinae and P. rubae. The parvus Subzone is readily correlated with sections at Among the Quadralella species, the angusta-dylani Subzone east Pardonet Hill and Juvavites Cove, where the interval is much sees the ascendency of subspecies of Q. praecommunisti and of Q. thinner (~70 cm thick at the former site). In all sections, the subzone willistonense, which can be distinguished by their pit position. Large is conspicuous in the dramatic decrease of typical Carnian conodont Q. noah and more ornate Q. kathleenae are less common through the elements referred to Kraussodontus, Parapetella, and Quadralella: subzone, and uncommon to rare Q. ex gr. oertlii, Q. posteroexpansa it represents a major faunal turnover (Orchard 2007c, fig. 5). Several subsp. A, Q. roysi, Q. sigmoidalis, and Q. tuvalica occur sporadically. species of Primatella also have their first appearance at the base of The angusta-dylani Subzone also contains additional Q. postlobata, the parvus Subzone but they are less common than the diminutive which is the probable origin of Q. pardoneti, as well as numerous small indices. Within the parvus Subzone, three further subdivisions can be elements assigned to Q. sp. indet. Also making a first appearance near differentiated based on the very rapid turnover of taxa. the base of the subzone and ranging through it are six Kraussodontus Lower subdivision (beds 18c-18e). taxa. Of these, K. roberti beta morphotype, K. rosiae, K. urbanae, and K. vancouverense are relatively common, K. margaretae, K. wendae, The lowest 40 cm of the parvus Subzone contains Metapolygnathus and K. sp. nov. A much less so. parvus plus the firstPrimatella asymmetrica, P. circulare, P. ovale, and Very few taxa are restricted to the angusta-dylani Subzone, and P. rhomboidale; these are joined slightly later by P. bifida. In addition, all are rare: Kraussodontus sp. nov. A, Parapetella aff. destinae, and Parapetella pumilio and P. aff. pumilio occur in a single sample P. rubae. The same is true of taxa that have their last occurrence: P. from this lower part but are not found higher. A single, specifically aff. asymmetrica, P. sp. nov. A and B, Q. pardoneti alpha morphotype, indeterminate specimen of Norigondolella also occurs. Several species and Q. stephanae. Long ranging taxa that appear earlier and range that first appear in the subjacent acuminata-prominens Subzone also throughout the subzone are also far less common than those which disappear by the end of this interval: P. posterolata, P. prominens newly appear. The former are Acuminatella sinuosa, Kraussodontus angulare, and Quadralella posteroexpansa subsp. B. Among the longer roberti alpha morphotype, Primatella mclearni, P. oblonga alpha ranging taxa that appeared earlier in the primitia Zone, 17 species morphotype, P. rectangulare, Quadralella postlobata, Q. roysi, and Q. disappear in this lower subdivision, whereas 9 new taxa appear (Fig. 6, tuvalica (Appendix, Table 4). Appendix, Tables 6, 7). Acuminatella acuminata-Parapetella prominens Subzone (beds 17- Middle subdivision (beds 18f-18h) 18b). The next 40 cm of the parvus Subzone contains the first appearance This subzone is equivalent to Fauna 6 of Orchard (2010b), and of several diminutive species: Parapetella johnpauli, P. willifordi, P. occurs in the highest ~1.3 m of the “Lower primitia Zone” sensu sp. nov. D, P. destinae beta morphotype, and Metapolygnathus parvus 33 gamma morphotype. In conjunction with these five first appearances, 18 E. aff. uniformis Orchard, E. aff. vialovi (Buryi), and E. sp. nov. A species disappear in the middle division, including all Kraussodontus appear concurrently. The “aff.” determination of these elements reflects species and virtually all Parapetella and Quadralella species (Fig. 6, the less ornate character of the original species from Europe and it is not Appendix, Tables 6, 7); only Acuminatella and Primatella species, clear whether any of those names should be used unequivocally. The the long ranging but uncommon Metapolygnathus ex gr. communisti, BBR taxa were formerly encompassed by the E. quadrata population and rare Parapetella species range higher. The two specimens of of Orchard (1983, fig. 4). Bed-by-bed sampling of the transition zone at Norigondolella recovered from these strata appear to be examples of BBR (Fig. 3, Appendix, Table 8) show the conodont change was abrupt, N. navicula, which does not make a further appearance until it becomes and none of the constituent species of the quadrata Zone appear earlier abundant above the boundary interval. than the others. The quadrata Zone extends through ~15 m of strata at Upper subdivision (beds 20a-21f). BBR but has not been studied in detail above the basal meter. The greater part of the parvus Subzone, about 2+ m thick at BBR, Epigondolella triangularis Zone. is assigned to the upper subdivision of the subzone. The top is drawn The succeeding triangularis Zone is characterized by more at the disappearance of Metapolygnathus parvus and all the associated strongly ornate platform elements that have not been studied in detail diminutive taxa. Within this upper division, the last Quadralella for this work. The occurrence of the triangularis Zone and that of elements disappear, but no new taxa appear. Associated Primatella younger Norian zones are summarized by Orchard et al. (2001b, c). species (except P. rhomboidale), plus two species of Acuminatella continue into the next and last subzone of the primitia Zone, which is INTEGRATED BIOSTRATIGRAPHY AT completely dominated by these ornate forms (Fig. 6, Appendix, Tables BLACK BEAR RIDGE 6-8). Orchard et al. (2001b, c) summarized the paleontology of the BBR section as a whole, which contains abundant ammonoids, bivalves, Primatella asymmetrica-Norigondolella Subzone (beds 21g-10/4). conodonts, and ichthyoliths indicating the Upper Carnian of the Upper This interval was referred to as the Metapolygnathus primitius Triassic through the Hettangian of the Lower Jurassic. A description of Zone by Orchard (2007c), Fauna 8 by Orchard (2010b), and the Upper the main faunal elements through the CNB interval is provided here primitia Zone by Orchard (2013). The subzone, which is ~15 m thick within the framework of the newly defined conodont zonation, and with at BBR (compared with ~4.5 m at PHE and 30+ m at Juvavites Cove), an emphasis on integration of the fossil faunas. Primary chronology is is defined as the range of Primatella species above the disappearance provided by the ammonoid zones of the Upper Carnian Klamathites of M. parvus and below the appearance of Epigondolella quadrata macrolobatus Zone, and of the Lower Norian Stikinoceras kerri and sensu stricto. The arbitrary choice of P. asymmetrica as a nominal Malayites dawsoni zones recognized at BBR (Fig. 3). This integration species is influenced by the fact that it is the most common of the few is supplemented through contemporaneous occurrences elsewhere on species restricted to stratigraphic levels above the base of the parvus the shores of Williston Lake, that is (from east to west) at McClay Spur, Subzone; that it occurs throughout the present subzone; and that it is east Carbon Creek, Brown Hill, Pardonet Hill east, and Juvavites Cove known to occur elsewhere in North America. As discussed above, the (Fig. 1). In all cases, undisputed Kerri Zone ammonoid collections asymmetrica-Norigondolella Subzone is not precisely the equivalent yield conodonts of both the asymmetrica-Norigondolella Subzone of of the “Upper primitia Zone” identified by Orchard (1983, 1991b), the primitia Zone and, in the uppermost Kerri Zone, conodonts of the which was differentiated on the first appearance of facies-dependent quadrata Zone (Orchard, 1983, 1991b). In contrast, the Upper Carnian Norigondolella. However, the presence of abundant Norigondolella Macrolobatus Zone is only known, apart from BBR, from pre-flood is only known from stratigraphic levels above the parvus Zone (Fig. localities on Pardonet Hill, and farther afield in British Columbia. 3, Appendix, Tables 6-8), and its presence in significant numbers with Intercalibration of the Upper Carnian ammonoid and conodont zones Primatella spp. may be used as a guide to this subzone. Furthermore, is facilitated by the use of matrix samples isolated from archive preliminary analysis of the large collections of Norigondolella from Macrolobatus Zone ammonoid collections (see below). this subzone permit the differentiation of N. norica sp. nov., which Few macrofossils occur in the Ludington Formation at BBR, but occurs only in beds 25 and 26 at BBR, and of two morphotypes of N. they become much more common in the Pardonet Formation. Among navicula that may also have stratigraphic significance. the first are brachiopods that occur in a distinctive shell bed near the The entire asymmetrica-Norigondolella Subzone is characterized base of the formation: this is a useful marker and its base was used by mostly long-ranging Primatella species, most of which first appear as datum for measuring the section (0 m, Figs. 2-4). The overlying in the angusta-dylani Subzone and only become dominant in the beds contain a succession of bivalves and ammonoids that have been upper subdivision of the parvus Subzone (Fig. 6). Most Primatella summarized recently by McRoberts (2011), McRoberts and Krystyn species occur throughout the subzone, and there are very few new (2011), and Balini et al. (2014). In addition, there are uncommon appearances that provide potential to subdivide the interval. Rare nautiloids, crinoid bioclasts, and belemnoids, and the acid-insoluble Acuminatella curvata appear in the lowest part of the subzone where microfossil residues yield common ichthyoliths, microbivalves, they are associated with the last occurrences of Primatella bifida microgastropods, echinoderm fragments, and uncommon cephalopod and P. rotunda. Above those strata, Norigondolella norica occurs in arm-hooks. The ichthyoliths were studied by Johns et al. (1997), and some abundance before being replaced by the alpha morphotype of N. summarized in Orchard et al. (2001b, c). Zonneveld et al. (2010a) navicula. Similarly, in the top 2 m of the subzone, rare Primatella sp. provided a thorough description of the sedimentary units and facies nov. A and distinctive morphotypes of Primatella ex gr. pseudoechinata of the BBR section, providing context for the biostratigraphic features occur during a brief re-appearance of Parapetella species, including P. identified. The occurrence of the macrofossils and the horizons sampled destinae and P. sp. nov. G. None of these taxa are common, but they for conodonts are shown on Figure 3; numbered beds are also conodont may prove useful for indicating the uppermost part of the subzone. samples, with bed numbers stabilized in 2001 et seq. The features of each conodont zone and subzone are given below. Epigondolella quadrata Zone (beds 10/5-32+). The samueli Zone at BBR is sparsely fossiliferous except in the The top of the studied lower section at BBR shows a rapid top 2 m of beds, which are low in the Pardonet Formation. These strata turnover from Primatella faunas to those dominated by Epigondolella (Z8, Z9 of Zonneveld et al., 2010a) have produced conodonts of the quadrata and affiliated forms (Fauna 9 of Orchard, 2010b). The base successive medioconstricta and spenceri subzones accompanied by of the quadrata Zone, defined by the appearance of the name-giver abundant and diverse actinopterygian (bony fish) teeth and ganoid (Orchard, 1991a, p. 179), marks the disappearance of virtually all scales, and elasmobranch (shark) scales and teeth including abundant Primatella species and their replacement by more strongly denticulate Labascicorona mediflexura Johns (Johns et al., 1997). Sedimentary homeomorphs. Norigondolella is also absent. Only Primatella destinae unit Z9 is the 60 cm thick marker bed containing abundant brachiopods and the small P. binodosa, which probably also includes juvenile concentrated in several layers: it is composed of monotypic elements of Epigondolella, survive into the quadrata Zone, although rhynchonellids referred to Piarorhynchia winnemae (Smith) (Sandy, the morphological spectrum of Epigondolella species that replaces in Orchard et al., 2001b, c.). This bed, which has so far yielded only those of Primatella often mimic them. uncommon indeterminate ammonoids and bivalves, is assigned to the In addition to abundant E. quadrata, common E. aff. miettoi spenceri Subzone of the samueli Zone. Mazza, Cau and Rigo, and less common E. aff. stefanoniensis Noyan, The lowest subzone of the primitia Zone, the sagittale-beattyi 34 Subzone, contains the ammonoid Thisbites sp. at its base, which (Krystyn, 2011). Also in bed 18d, the ammonoid Tropiceltites occurs implies these strata are no older than Macrolobatus Zone (Tozer, 1994). with Pterosirenites and Griesbachites, and in beds 18f-g, further The first of two levels for the ammonoid Anatropites sp. occurs slightly examples of Pterosirenites and Gonionotites occur with the same higher (Tozer, in Orchard et al., 2001b, c) in Unit Z10 of Zonneveld Halobia species. None of these elements are diagnostic for a particular et al. (2010a), and this is now supplemented with a Tropiceltites ammonoid zone, although the appearance of species of Pterosirenites is from an equivalent level (bed 8; pers. commun., L. Krystyn, 2010). taken as defining the base of the Norian in the Boreal Realm (Krystyn, Anatropites is the most important taxon for the identification of the 2011; Balini et al., 2012), so the association of H. austriaca and Macrolobatus Zone (Fig. 3) and confirms that the base of the primitia Pterosirenites in these beds suggests a Norian age (Balini et al., 2012). Zone, as redefined, corresponds closely to the base of this uppermost The above occurrences all lie within the lower and middle subdivisions Carnian ammonoid zone. The bivalves Halobia septentrionalis of the parvus Subzone, whereas the stratigraphically thicker upper and H. ornatissima (McRoberts, 2011) also make an appearance in subdivision of the subzone contains mostly bivalves and lacks new the sagittale-beattyi Subzone. A change in the ichthyolith faunas taxa. Of note, Halobia radiata last occurs in this upper subdivision introduces the Synechodus multinodosus concurrent-range ichthyolith (bed 21c). The parvus Subzone marks a major turnover in the conodont Zone (Johns et al., 1997, in Orchard et al., 2001b, c), for which BBR is fauna, and virtually all inornate species typical of the Carnian disappear the type locality. This zone is marked by a peak abundance of the latter by the close of the middle subdivision (Fig. 6, Appendix, Tables 6, 7). elasmobranch tooth as well as Minuticorona scales, actinopterygian After M. parvus and its diminutive associates disappear, and 0.5 m teeth and scales, and other ichthyoliths. above the base of the overlying asymmetrica-Norigondolella Subzone, The succeeding angusta-dylani Subzone of the primitia Zone the ammonoid Guembelites clavatus (McLearn) occurs (in bed 22b, broadly corresponds to sedimentary unit Z12 of Zonneveld et al. Fig. 28). This ammonoid is diagnostic for the lower subzone of the (2010a). It includes ammonoid occurrences both near its base (bed Kerri subzone I (Tozer, 1994, p. 39). About 15 m above Guembelites 13) and near its top (beds 16-17), the latter being a second level for in bed 31 (10/3; Fig. 3), subzone II of the Kerri Zone is recognized by Anatropites, indicating the Macrolobatus Zone. As noted by Tozer the occurrence of Stikinoceras kerri McLearn. This occurrence is very (1994, p. 38), the faunas of the Macrolobatus Zone differ one from close to the top of the asymmetrica-Norigondolella Subzone, and of the another and probably are not the same age. The truth of this statement is Primatella primitia Zone. Hence, the primitia Zone extends no higher obvious at BBR where at least two conodont subzones are identified in than the lower part of Kerri subzone II. The succeeding Bed 32 and sub- Anatropites-bearing beds, and further age variation becomes evident as beds 10/5-7 contain conodonts of the Epigondolella quadrata Zone. one considers conodont data from other Macrolobatus Zone localities Bivalve species disappear in a staggered fashion during the (see below). Among the bivalves, the same species of Halobia present asymmetrica-Norigondolella Subzone. Halobia ornatissima ranges in the preceding subzone are joined by H. radiata (McRoberts, 2011), up through the lower one-third of the subzone, H. selwyni extends which makes its first appearance at the base of this subzone. Johns (in up to about midpoint of the subzone, and H. austriaca continues on Orchard et al., 2001b, c) reported a mixture of ichthyoliths from this alone (McRoberts, 2011, fig. 9). Additional halobiids that occur in subzone, including samples in which deeper water actinopterygian the overlying quadrata and triangularis zones are H. beyrichi and H. scales and teeth were common to abundant but no elasmobranch teeth cordillerana (Orchard et al., 2001b, c; McRoberts, 2011, fig. 8). or scales were recovered. The top of subzone II of the Kerri Zone occurs 5 m higher than The overlying acuminata-prominens Subzone marks the beginning its base and is quickly followed by three levels of the Dawsoni Zone of the CNB interval sampled in greater detail (Figs. 27, 28). It contains the ammonoids, starting about 40 m above the section datum and extending ammonoid Gonionotites and further examples of the bivalves Halobia over some 8.3 m (Fig. 3). Within the Dawsoni Zone, the conodont septentrionalis and the long ranging H. ornatissima. Conodont yield in faunas change from those of the quadrata to the younger triangularis this subzone is very high (Fig. 6), implying slowdown in sedimentation Zone, but that interval has yet to be studied in detail. The triangularis and/or an increased mortality of the conodont animals. Notably, the Zone contains the first new elasmobranch ichthyoliths typical of the entire boundary interval - including this subzone, the overlying parvus Norian (Johns et al., 1997), including Synechodus incrementum Johns, Subzone, and lowest part of the asymmetrica-Norigondolella Subzone- Fragilicorona, and Glabrisubcorona, an assemblage that extends into are developed within a single facies association of thin- to medium- the Middle Norian. bedded turbidites of a single stratigraphic unit (Fig. 27; Zonneveld et al., 2010a, FA2, Z13). Complementary data for the CNB placement in British Columbia Within the next higher parvus Subzone, there is evidence of faunal Apart from the co-occurrences of macrofossils and conodonts turnover in several groups. At the base of the subzone, in bed 18c, the documented at BBR, there are important boundary sections at nearby bivalve Halobia septentrionalis has its last occurrence. In the following Pardonet Hill east (PHE) and Juvavites Cove. Several other sections bed, 18d, Halobia selwyni and a questionable H. austriaca (or juvenile include Kerri Zone faunas, but the underlying strata provide few of another species?) appear (Fig. 28; McRoberts, 2011, pers. commun.). additional data. In addition, seven conodont collections have been The latter species, which definitely occurs 30 cm higher in bed 18f, recovered from the matrix of Macrolobatus Zone ammonoids from is reported to appear at the base of the Guembelites jandianus Zone western Canada, four from northeastern British Columbia, and three in Austria, regarded as equivalent to the North American Kerri Zone from the far west on Haida Gwaii, in the allochthonous Wrangel Terrane

Figure 27. Photograph of the CNB beds in the lower Black Bear Ridge section (detail of Figure 26). Bed numbers correspond to those shown in other figures. Subzonal boundaries of theprimitia Zone and subdivisions of the parvus Subzone are indicated. 35

Figure 28. The CNB interval at Black Bear Ridge showing (from left) sample numbers, macrofossil (bivalve, ammonoid) occurrences and ranges (gray dots), conodont subzones, and key conodont ranges (black dots) for the ~5 m interval (see Appendix for all occurrences). Conodont genera abbreviations as in Figure 4. 36 (Orchard, 1991a). identified in the basal 10 cm of the exposed Pardonet Formation, which CNB beds at the PHE section (Figs. 29, 30) include a well exposed is otherwise unfossiliferous. Hence, some part of the parvus Subzone outcrop showing the contact between the Baldonnel and Pardonet and three subjacent subzones of the primitia Zone are not preserved at formations. This Baldonnel section represents a shallower water Upper the contact. This hiatus is greater than that recognized at PHE, although Carnian environment than that at BBR, where the Pardonet overlies it may be compounded by minor fault movement at the formational the Ludington Formation. Pardonet Hill east lies close to the hinge boundary. line separating the two Carnian formations. The Baldonnel Formation Ammonoids and conodonts are associated in two archival has few macrofossils near its top (see Zonneveld et al., 2007 for an Macrolobatus Zone collections made by E.T. Tozer in 1964 from account of older faunas), but they become common in the Pardonet the Pardonet Formation of pre-flood Pardonet Hill, whose probable Formation, the lowest 2 m of which has been collected bed-by-bed. location is close to the present Juvavites Cove. These collections Both Pterosirenites and Halobia austriaca occur in the basal bed are GSC [Geological Survey of Canada] curation numbers O-64616 of the Pardonet Formation and over the following meter of section and, from 0.6 m above, O-64628 (Tozer, 1994, p. 323; Fig. 30). The (Tozer, pers. commun., 2001; Krystyn, pers. commun., Dec. 2009; former sample yielded the key conodonts Acuminatella acuminata McRoberts, pers. commun., July 2013). Conodonts recovered from the and Primatella asymmetrica plus an array of other pre-turnover taxa, basal bed and “transitional strata” immediately beneath it belong to the but it lacks Metapolygnathus parvus: its conodont age is regarded as acuminata-prominens Subzone, and those from the following 70 cm close to the boundary between the acuminata-prominens and parvus represent the parvus Subzone (Fig. 29). This succession is comparable subzones (Fig. 31). The younger archive collection O-64628 yielded to that at BBR except that the firstPterosirenites and Halobia austriaca all morphotypes of M. parvus in addition to Parapetella destinae alpha are slightly older at PHE. In common with BBR, Guembelites appears morphotype, P. prominens, and P. pumilio, and many additional taxa within the overlying asymmetrica-Norigondolella Subzone, as do that disappear at BBR by the end of the middle subdivision of the abundant Norigondolella. parvus Subzone, to which interval the collection is assigned (Fig. 31). Conodonts indicative of the samueli Zone occur at the top of the The character of these archival collections has far-reaching massive Baldonnel Formation at PHE and imply a hiatus has cut out at implications. Although they represent stratigraphic record not seen at least the lower subzones of the primitia Zone that are present at BBR. Juvavites Cove at present, they can be projected to fall directly beneath This is supported by the occurrence of a bone bed at the boundary. the fauna that is preserved there in the Pardonet Formation. As such, Reduced sedimentation is also reflected in the total thickness of the they complement the preserved stratigraphy and make it comparable parvus Subzone, which is about one-quarter of that at BBR, and in with that preserved at PHE. More importantly, the association the relative condensation of the lower and middle subdivisions of the demonstrates that the Macrolobatus Zone extends through the subzone. Most of the CNB interval is taken up by the upper subdivision acuminata-prominens Subzone and as high as the middle subdivision of the parvus Subzone. Common Norigondolella appear about 0.5 m of the parvus Subzone. This cannot be demonstrated at BBR, where above the parvus Subzone at PHE, but about 4 m above that datum at the highest Anatropites occurs at the top of the subjacent sagittalis- BBR, in both cases after the appearance of the ammonoid Guembelites. dylani Subzone. Furthermore, it implies that the parvus Zone with Finally, the occurrence of Stikinoceras, indicative of the upper Kerri Pterosirenites and Halobia austriaca at both BBR and PHE are time Zone, is 14 m above the top of the parvus Subzone at BBR, but about 1 equivalent to the Macrolobatus Zone and are, therefore, of Carnian (in m above that level at PHE. the traditional sense) rather than the Norian age that has been favored About 1 km to the west from PHE section, a less well exposed (see above). This conclusion identifies two anomalies: the absence of boundary section occurs at Juvavites Cove (Fig. 30). In common with the genus Anatropites in the parvus Subzone of BBR, and the absence of the former section, the uppermost beds of the Baldonnel Formation Pterosirenites in the archival Macrolobatus collections from Pardonet yield samueli Zone conodont faunas, in this case specifically the Hill. There is no obvious explanation for this other than collection spenceri Subzone. This Upper Carnian fauna was recovered adjacent failure, or perhaps differing ammonoid habitats. to a poorly exposed breccia in the narrow creek bed and co-occur with The conodont zonal assignments of all Macrolobatus Zone brachiopods, as they do at BBR. In the juxtaposed cliff exposure of collections are summarized in Figure 31: clearly, the ammonoid zone the lower Pardonet Formation, the first ammonoids occur about 2 m extends over several conodont subzones. The oldest record is from above the exposed base (Fig. 30): these include ammonoid indices for the beattyi-sagittalis Subzone at BBR, although this is based only on the lower Kerri Zone, that is, Discostyrites ireneanus and Guembelites Thisbites. The second level is high in the angusta-dylani Subzone at clavatus. As in all other sections, this ammonoid fauna co-occurs with BBR, which correlates with an additional archival collection from conodonts of the asymmetrica-Norigondolella Subzone of the primitia Mount McLearn (GSC cur. no. O-068202; Tozer, 1994, p. 328); both Zone. Underlying this, the upper subdivision of the parvus Subzone is these collections are ~7.5m below Kerri 1 ammonoids. The Mount

Figure 29. Photograph of the CNB section at Pardonet Hill east (see Figure 1) showing bed numbers and conodont zones and subzones. The top of the Baldonnel Formation on the left belongs to the samueli Zone and is overlain disconformably by bone beds referred to the acuminata-prominens Subzone at the base of the Pardonet Formation. Macrofossil occurrences are shown in Figure 30. 37

Figure 30. Conodont based correlation between the CNB interval at Black Bear Ridge, Pardonet Hill east, and Juvavites Cove (Figure 1), with macrofossil occurrences shown in context of conodont subzones. Thick black line at 0 m of latter two sections indicates hiatus below Pardonet Formation. The relative position - in terms of their conodont content - of two archival ammonoid collections from Pardonet Hill are shown on the lower right. Also indicated are the stratigraphic separation of the Norigondolella blooms (Ng), the conodont quadrata Zone, and the ammonoid Dawsoni Zone (measured from the 0 m datums in each section). 38

Figure 31. Composition of Macrolobatus Zone ammonoid faunas (largely after Tozer, 1994), accompanying halobiids on Huxley Island (C. McRoberts, pers. commun., April 2014), and their assignment to conodont subzones of the primitia Zone. The oldest collections are in section at Black Bear Ridge, two are archive from Pardonet Hill (see Figure 30), two are archive from elsewhere in northeastern B.C. (Mount Laurier, Mount McLearn), and three are from Haida Gwaii on the west coast (Kunghit and Huxley islands). This demonstrates that the stratigraphic scope of the ammonoid zone embraces the entire primitia Zone at Black Bear Ridge, including the ~5 m CNB interval that lacks ammonoid zonal indices. The position of the lower and upper Kerri Zone indices (vertical bars) at BBR is also shown. McLearn collection includes Parapetella columbiense, P. cordillerense, Macrolobatus Zone extends from near the base of the primitia Zone and P. rosiae in the absence of Acuminatella acuminata. A fourth through four conodont subzones up to the basal beds of the fifth, the archival collection, from Mount Laurier (O-94738; Tozer, 1994, p. P. asymmetrica-Norigondolella Subzone, which is very close to the 341), is also assigned to the angusta-dylani Subzone based on the first occurrence of Guembelites. Hence, the first occurrences of both presence of Acuminatella sagittale, A. angusta, and additional diverse Pterosirenites and Halobia austriaca lie within the Macrolobatus Zone. taxa. The Macrolobatus collection from the Peril Formation on Kunghit Island on Haida Gwaii (C-157382; Tozer, 1994, p. 348) comes from Correlation with Pizzo Mondello, Sicily the historical type location of Acrochordiceras carlottense (Whiteaves) Correlation between the GSSP candidate sections at BBR and collected by G.M. Dawson in 1878. The conodont fauna is sparse but Pizzo Mondello (PM) in Sicily based on conodonts has been hampered is apparently also assignable to the angusta-dylani Subzone based on by both differing faunas and differing taxonomic approaches. This Quadralella praecommunisti and the absence of younger indices. work on conodonts from BBR introduces substantial nomenclatural Two additional Macrolobatus collections from Huxley Island change at the generic level, and many newly differentiated species and on Haida Gwaii appear to extend the scope of that ammonoid zone morphotypes. Some of these taxa can be recognized in the published into even younger stratigraphic levels, and are notable for containing illustrations of PM conodonts (Mazza et al., 2010, 2011, 2012a, b), Halobia austriaca (C. McRoberts, pers. commun., April 2014): and a re-evaluation of that fauna may eventually add to this record. these collections are assigned GSC curation numbers C-157119 and Certainly there are differences, including endemic species in both C-157123. Although separated from one another by a small fault and regions. In comparing the conodonts, the following nomenclatural having different ammonoid faunas, these two collections were thought differences are noted between the present work and that of Mazza et al. by Tozer (1994, p. 348) to probably originate from the same level. The (op. cit.): first, structurally isolated and smaller collection (C-157119) contains Carnepigondolella sensu Mazza includes many species common Metapolygnathus parvus and lacks pre-turnover taxa. The second (C- to both areas. Some are assigned to that genus, e.g., C. anitae, C. 157123) is dominated by Primatella species, including P. circulare, pseudodiebeli, C. samueli, and C. zoae, but others are here included and sits about 1 m above similar Primatella faunas that include M. in Quadralella, e.g., Q. angulata, Q. carpathica, and Q. tuvalica, and parvus and are therefore comparable with the first collection, regarded exceptionally in Primatella, e.g., P. orchardi. as dating from the upper subdivision of the parvus Subzone. The Paragondolella sensu Mazza also includes taxa included in second collection falls within the basal part of the P. asymmetrica- Quadralella by this author, e.g., Q. noah and Q. oertlii. Few or no Norigondolella Subzone. Carnian species are included in Paragondolella by the present author. In summary, it can be demonstrated that the Upper Carnian Within Metapolygnathus spp., M. communisti and its derivatives, 39 including M. parvus, are retained in that genus, but M. praecommunisti others here assigned to Kraussodontus and Parapetella, which become is re-assigned to Quadralella, and M. mersinensis and M. cf. primitius more common at this level and include many platform elements with are re-assigned to Primatella. Among elements assigned to Q. anteriorly shifted pits, taken by Mazza et al. (2012b) as diagnostic of praecommunisti at PM, the author recognizes Parapetella clareae and Metapolygnathus. These elements appear low in the primitia Zone at Kraussodontus roberti. BBR, after the extinction of Carnepigondolella at the top of the samueli The concept of Epigondolella sensu Mazza (2012b) is broader than Zone. Hence, it is probable that T2 approximates the samueli-primitia that applied in this work and includes several Tuvalian species referred zonal boundary (Fig. 5). to E. vialovi, E. heinzi, E. miettoi, and E. quadrata (intermediate The third turnover at PM, T3, occurs immediately after the form). These are much older than E. quadrata sensu stricto, which “boundary interval” of Mazza et al. (2012b) where Metapolygnathus is regarded as the oldest Epigondolella species in British Columbia. largely disappears and is replaced with “advanced” Epigondolella Younger Epigondolella species present in the basal Norian have been plus Norigondolella (between samples NA41 and PM30A). A parallel recorded by Mazza et al. (2012b) as E. praetriangularis, E. quadrata can be drawn with BBR in so much as Norigondolella only becomes intermediate forms, E. rigoi, E. spatulata, E. triangularis, E. uniformis, common in the asymmetrica-Norigondolella Subzone of the primitia and E. vialovi. Many of these species have holotypes from much Zone in faunas above the disappearance of most Carnian taxa, younger Norian faunas, and their occurrence at that level is similarly exemplified byMetapolygnathus sensu Mazza et al. (2012b). In British anomalous. The appearance of “advanced forms” of Epigondolella Columbia, rare Metapolygnathus and Parapetella species continue quadrata some 45 m higher at PM would, in this author’s view, more through the primitia Zone but the faunas are dominated by Primatella accurately align with the base of the quadrata Zone at the top of the and Norigondolella (Figs. 3, 6). The “advanced” Epigondolella species BBR succession. The older PM material should be re-assessed as referred to by Mazza et al. (2012b) may be examples of Primatella, possible representatives of Carnepigondolella or Primatella of this although, as discussed above, many of the holotypes of the identified report. species in this PM interval were originally described from younger Primatella currently accommodates the Sicilian species assigned Norian faunas. The first true examples ofEpigondolella appear near the to Carnepigondolella orchardi, C. pseudoechinata, Metapolygnathus top of the studied section at BBR, at the base of the “type” quadrata cf. primitia (= M. mazzai Karadi, Kozur and Gorog), and some elements Zone. This may correlate with the level just below the slump breccia of M. mersinensis. Examples of Primatella subquadrata (identified as where Mazza et al. (2012b, fig. 2) shows the appearance of advanced Carnepigondolella pseudoechinata and Metapolygnathus mersinensis forms of E. quadrata. This datum is high in the Lower Norian Kerri by Mazza et al., 2012b) occur in samples ~10 m below boundary Zone and is not the same datum (T1) advocated as a potential stage interval and imply the primitia Zone. C. gulloae is an endemic that may boundary by Mazza et al. (2010, p. 131). also belong to Primatella. In summary, the lower two of the three conodont bioevents Norigondolella occurs in both regions but includes the endemic identified at PM by Mazza et al. (2010) are well below a suitable species N. trinacriae at PM, and perhaps N. norica at BBR. Among the position for the CNB but may be equivalent to positions within, and at other new genera from British Columbia, Acuminatella is apparently the top of the samueli Zone. Level T3 is regarded as close to the base of absent from Sicily, whereas some Kraussodontus and Parapetella the asymmetrica-Norigondolella Subzone of the primitia Zone (Fig. 5). species are recognized. Acuminatella does occur in the quasi-Tethyan More recent consideration of the PM successions has identified Wrangell Terrane of the Pacific margin (Haida Gwaii; Desrochers three new conodont bioevents (Mazza et al., 2012b, p. 91) that all lie and Orchard, 1991; Carter and Orchard, 2013), and also in the more between events T2 and T3, that is bounded by the “boundary interval” of southerly latitudes of North America, in Nevada (Balini et al., 2014). Mazza et al. (2012b). These three datums are marked by, successively, With these nomenclatural differences in mind, comparison is the first occurrence of Metapolygnathus parvus, of “M. echinatus attempted with three major turnovers of conodont genera recognized sensu Orchard” = Parapetella destinae, and of Carnepigondolella = by Mazza et al. (2010) at PM. In their terminology, these events are: Primatella gulloae. Although the latter species does not occur at BBR, T1 - Carnepigondolella is replaced by Epigondolella (sample these PM datums can be respectively correlated with BBR at the base FNP88a), subsequent to the disappearance of Paragondolella (NA25). of the parvus Subzone, a position within the lower subdivision of the T2 - Metapolygnathus becomes abruptly dominant over parvus Subzone for P. destinae, and a position near the top of the upper Epigondolella (between samples AM23 and NA35). subdivision of the parvus Subzone for P. gulloae. The large turnover in T3 - Metapolygnathus largely disappears and is replaced with the Canadian section begins below the base of the parvus Subzone, and “advanced” Epigondolella plus Norigondolella (between samples continues through the entire span of the subzone up to an equivalent NA41 and PM30A). level to T3 at PM. The bloom of diminutive elements like M. parvus T1, the replacement of Carnepigondolella by Epigondolella, does and P. destinae in the absence of other Carnian holdovers characterizes not obviously occur at BBR where Carnepigondolella disappears at the the base of the upper subdivision of the parvus Subzone at BBR (Fig. top of the Carnian samueli Zone, and Epigondolella sensu stricto appears 6), but no such event is identified at PM. much later, above the succeeding primitia Zone. Carnepigondolella Metapolygnathus parvus is recorded over ~12 m at PM (Mazza species of Mazza et al. (2010) range higher than Carnepigondolella et al., 2012b, fig. 2) compared with ~3.5 m at BBR. In Sicily, itis sensu stricto at BBR and disappear at various levels within the primitia accompanied by M. communisti, which becomes very abundant at that Zone. This is also true for species included in Paragondolella by Mazza time (Mazza et al., 2012b, p. 91) and could conceivably be confused et al. (2010), which are mostly referred to Quadralella here, and which with the former. In contrast, M. ex gr. communisti is uncommon and allegedly disappear prior to T1. Even allowing for nomenclatural occurs only sporadically through the BBR section: it provides no guide differences, there is no obvious alignment in the disappearance of these to the boundary interval. However, within the elements assigned to various taxa in PM and BBR. M. communisti by Mazza et al. (2012b), there is one specimen here Regarding the replacing fauna of Epigondolella in the Tuvalian regarded as an example of Parapetella irwini. This specimen occurs (Mazza et al., 2012b), as outlined above, this is a much earlier appearance in sample NA37, within the boundary interval at PM, and from the for the genus than at BBR and represents a major contrast between the acuminata-prominens Subzone through the middle subdivision of two regions. Carnepigondolella pseudodiebeli beta morphotype, which the parvus Subzone at BBR. Another species that appears useful for appears in the eozoae-ludingtonensis Subzone of the samueli Zone, trans-Panthalassa correlation is Primatella bifida of this report (= is a homeomorph of E. quadrata: it appears below the extinction of Metapolygnathus linguiformis sensu Mazza et al., 2012b), which Carnepigondolella sensu stricto, as do the “Epigondolella” species from seems to align around the middle subdivision of the parvus Subzone. PM. One interpretation of T1 might align it with a position within the Dominance of the faunas by Primatella and Norigondolella occurs samueli Zone where Tethyan diversification of the Carnepigondolella in the asymmetrica-Norigondolella Subzone following the parvus stock may have occurred (Fig. 5). Subzone at BBR, and a little above the “boundary interval” at PM At T2, Mazza et al. (2010, p. 131) record Metapolygnathus becoming (where M. parvus overlaps with P. gulloae). dominant over Epigondolella, and the disappearance of the last of the Our knowledge of PM has also improved with modern studies “typical” Carnepigondolella species, C. pseudodiebeli; both C. gulloae on the ammonoids (Balini et al., 2012) and halobiid bivalves (Levera, and C. pseudoechinata occur later, but these are more appropriately 2012). Two ammonoid zones are recognized at PM, the Upper Carnian assigned to Primatella. This level may correspond to the disappearance Spinosus Zone and the Lower Norian Jandianus Zone. Between the of Carnepigondolella at BBR. For Mazza et al. (2010, 2012b), the two, there are 18 m of strata without diagnostic ammonoids (Balini et “replacement” fauna of Metapolygnathus can be re-interpreted as al., 2012, p. 79). This embraces the entire “boundary interval” at PM. the ascendency of Primatella spp., Quadralella praecommunisti, and The bivalve Halobia austriaca first occurs within this interval (sample 40 FNP135A), a short distance below Norian ammonoids and equivalent species may provide a proxy for this datum. Significantly, this level is to the position of turnover T3 (Balini et al., 2012, fig.12). now known to be equivalent to the boundary between the Macrolobatus Concerning chemostratigraphy, the small negative excursion of and Kerri zones, and of the traditional ammonoid-based CNB. 1.12‰ in δ13Corg occurring at (what can now be referred to as) the The appearance of Epigondolella quadrata clearly lies in the upper base of the middle subdivision of the parvus Subzone (Williford et al., Kerri Zone, not far beneath the succeeding Dawsoni Zone. It is far too 2007) was interpreted to be the result of a decline in net primary produc­ high to serve as a CNB index, just as the appearance of quadrata-like tion, and a potential cause of increased selective pressure on conodonts homeomorphs in the samueli Zone are far too low. and bivalves manifest as a significant turnover observed at this level. In contrast, a positive isotope shift of similar magnitude (1.2‰) based SUMMARY on carbonate was found in samples PM24-25 at PM, which lie lower Abundant and diverse conodonts from the Carnian-Norian down, beneath the boundary interval (Muttoni et al., 2004; Muttoni et Boundary (CNB) interval at the Global Stratigraphic Section and al., 2014). This was thought to be related to transient paleogeographical Point (GSSP) candidate section at Black Bear Ridge (BBR), British or paleoenvironmental conditions in the area that are also reflected Columbia, are systematically described and re-classified with emphasis in the conodont turnover characterized as T2, here interpreted to lie on the profile and sculptural features of the anterior platform margins, well below the boundary interval at BBR. Whatever the cause of these posterior carina development, posterior platform shape and ornament, different isotopic signatures, they provide no basis for correlating the and position of the basal pit. This results in a substantially new taxonomy two sections. for the interval. Based on over ~35,000 conodont elements recovered Conodont Datums for the CNB from about 250 kg of carbonate rock, five new genera (Acuminatella, Kraussodontus, Parapetella, Primatella, Quadralella), 111 species (75 By 2007, the CNB interval between proven Macrolobatus and Kerri newly described here), six new subspecies, and 47 new morphotypes Zone faunas at BBR was reduced to <5 m of strata between beds 17 and are differentiated. This doubles the number of platform genera, and 22 (Fig. 3). Within this interval, a significant conodont faunal turnover increases by a factor of three or four the number of taxa known from was identified between beds 18 and 20. Bed 18 and underlying strata the interval compared with the most recent data from PM (four genera, produced a rich and diverse conodont fauna dominated by the typical 46 lesser taxa: Mazza et al., 2012b, fig. 2). Carnian genera Kraussodontus, Quadralella, and Parapetella, with a Proposed phylogenies of the BBR conodont taxa are shown in minor percentage of Acuminatella and Primatella (Fig. 6). In contrast, 19 figures illustrating range and hypothetical relationships. Novel bed 20 produced a fauna dominated by Primatella accompanied by ancestor-descendant linkages are proposed for Carnepigondolella with many diminutive “survivor” elements, including Metapolygnathus diversification entirely restricted to thesamueli Zone, and extinction of parvus, Parapetella destinae, P. johnpauli, P. pumilio, and P. willifordi; the genus at the top of that zone. Acuminatella, which may be a North these species apparently represent the final chapter in several lineages of American endemic, and Primatella appear concurrently and serve as Carnian conodont genera. Primatella was seemingly unaffected in this useful guide fossils. These ornate species have been confused with both interval and flourished as the primary component of most subsequent Carnepigondolella and Epigondolella, but they are here regarded as the conodont faunas prior to the appearance of Epigondolella. Floods of predominant genera in the basal Norian prior to their demise concurrent Norigondolella elements starting in bed 25 (Fig. 3) provide a Norian with the later appearance of true Epigondolella in the quadrata Zone. signature. Metapolygnathus, used in a restricted sense, includes the long-ranging In the bed-by-bed resampling of the 5 m of strata between beds communisti group that gave rise to the diminutive M. parvus in the 17 and 22, additional macrofauna was collected, and samples were CNB interval. Parapetella is recognized as an important new genus that taken for carbon isotope analysis. Abundant conodonts were recovered evolves rapidly through the Upper Carnian along several lines, some of from each bed (Fig. 6). The results of this multidisciplinary approach which culminate also in diminutive elements in the boundary interval. were significant inasmuch as paleontological and chemical events were Quadralella and Kraussodontus are more conservative elements (many discovered to be well aligned: Pterosirenites sp., a potential Norian previously assigned to Paragondolella) that often dominate Carnian indicator, was collected over 0.5 m; the last Halobia septentrionalis was collections but which disappear during the big turnover in the CNB superseded by H. austriaca, a second potential Norian indicator, within interval. the same 0.5 m; and a small but distinct carbon negative excursion The conodont taxa form the basis for the definition of two also fell within this narrow interval, represented by beds 18e and 18f. conodont zones and nine constituent subzones for the interval. These The first appearance of Metapolygnathus parvus in bed 18c heralded are, in ascending stratigraphic order, the Carnepigondolella samueli both a sharp decline of typical Carnian taxa and rapid introduction of Zone with the subzones of C. eozoae-Kraussodontus ludingtonensis, diminutive “end-members” of these lineages. This facilitated further C. zoae, C. medioconstricta, and C. spenceri, followed by the subdivision of that parvus Subzone into lower, middle, and upper parts. Primatella primitia Zone with the subzones of Acuminatella sagittale- The Middle subdivision corresponds to the appearance of diminutive Parapetella beattyi, A. angusta-Metapolygnathus dylani, A. acuminata- Parapetella johnpauli and P. willifordi, the peak isotope minimum, Pa. prominens, M. parvus (three subdivisions), and the Primatella and the first undisputed Halobia austriaca. The Upper subdivision and asymmetrica-Norigondolella sp. These strata are capped by the Lower greater part of the parvus Subzone marked the virtual completion of Norian Epigondolella quadrata Zone of Orchard (1991b). the faunal turnover with the disappearance of most Carnian taxa and no Conodont faunal turnovers are identified at the boundary between further additions to the conodont fauna. the samueli and primitia Zones, between the primitia and quadrata Conodont datums within the boundary interval are the base of zones, and especially within the parvus Subzone. In each case, these the acuminata-prominens Subzone, the base of the parvus Subzone turnovers are marked by high conodont yield. The turnover that begins and of each of its subdivisions, and the base of the asymmetrica- just before the parvus Subzone and ends at the top of that subzone sees Norigondolella Subzone. As discussed above, several of these are the disappearance of 63 taxa belonging to four genera. This turnover is potentially recognizable at PM. The appearance of Metapolygnathus most pronounced over a ~2 m interval at BBR that coincides with the parvus (base bed 18c) occurs before the main demise of the Carnian appearance of Halobia austriaca, the ammonoid Pterosirenites, and a taxa at BBR, but the base of this subzone can be delineated at PM. carbon isotope minimum; both bioevents have been favored as Norian The base of the middle subdivision of the parvus Subzone (base of bed rather than Carnian indicators. However, no definitive ammonoids 18f) has the complementary first appearance of Halobia austriaca, and occur in the ~5 m CNB interval that is bracketed by Upper Carnian the isotope minimum. This level, based on the First Appearance Datum ammonoids of the Macrolobatus Zone, and those of the Lower Norian (FAD) of the bivalve, has been favored as a GSSP datum by McRoberts Kerri Zone. These “undated” strata are assigned to the acuminata- and Krystyn (2011), but, as discussed earlier, H. austriaca appears at prominens Subzone through the parvus Subzone of the primitia Zone. a lower stratigraphic level at PHE. This FAD datum can be delineated Matrix of archival ammonoid collections of the Macrolobatus by the appearance of Parapetella destinae at PM. The base of the upper Zone from British Columbia yield conodonts that can be assigned to the subdivision (base of bed 20a) is easily recognized at BBR and at other angusta-dylani Subzone, acuminata-prominens Subzone, and several localities nearby based on its relatively low diversity conodont fauna, levels within and just above the parvus Subzone, all within the primitia but it is not known whether this event can be recognized at PM. Finally, Zone. These complement the Macrolobatus collections from BBR, the disappearance of M. parvus and its diminutive associates at the top which occur in the even older beattyi-sagittale Subzone, and at the of the parvus Subzone is a boundary option that is broadly recognizable top of the succeeding angusta-dylani Subzone. The younger conodont at PM as T3. This datum is, however, largely based on disappearance of subzones identified in association with Macrolobatus ammonoids taxa rather than first appearance, although some uncommonPrimatella occur in the 5 m ammonoid gap at BBR and therefore imply that the 41 Macrolobatus Zone embraces most of the boundary interval prior to SYSTEMATIC PALEONTOLOGY the asymmetrica-Norigondolella Subzone and the first occurrence of Foreword Guembelites, diagnostic of the lower Kerri Zone. Other sections complement the data from BBR. At Pardonet Hill The following is a form taxonomy based on pectiniform P1 east (PHE), topmost acuminata-prominens Subzone, several levels elements. A multielement approach is precluded due to insufficient within the parvus Subzone, and the lower part of the asymmetrica- representation of the ramiform elements that are assumed to have Norigondolella Subzone are identified in strata containing both formed part of the original conodont apparati. Most of the taxa Pterosirenites and Halobia austriaca. Most of these collections are are ozarkodinid platform genera belonging to a single family, the now known to be correlatives of the Macrolobatus Zone. Collections Gondolellidae Lindstrom 1970, and are described in alphabetical order; from Juvavites Cove (JC) are complementary in containing parvus non-platform elements of uncertain familial position (Misikella) follow. Subzone conodonts beneath Kerri Zone ammonoids, and two archival Unless stated otherwise, the origin of all illustrated specimens, and the Macrolobatus Zone collections from nearby correlate with the type locality for all new taxa, is Black Bear Ridge, northeastern British lower beds of the Pardonet Formation at PHE. Additional archive Columbia, Canada. Macrolobatus Zone collections from further afield in Haida Gwaii also Total conodont element counts for each sample are given in the contain Halobia austriaca, confirming its appearance in the Carnian. Appendix Tables 1 and 2, which also provide the curation number At both PHE and JC, a hiatus occurs at the base of the Pardonet (formerly location number, prefixed with “C-”, “O-”, or “V-”). Formation at the boundary with the underlying Baldonnel Formation. Specimen counts for each species are summarized under “Stratigraphic In each section, the lowest part of the primitia Zone is missing: this Occurrence” as rare (1-5 specimens), uncommon (6-10), common (11- is equivalent to about 15 m stratigraphic thickness at BBR within the 30), very common (31-50), or abundant (>50 specimens). All illustrated relatively expanded, deeper water succession. Thereafter, the rate of material is deposited in the National Type Collections of the Geological sedimentation in the boundary interval (acuminata-prominens through Survey of Canada and bear type numbers prefixed with “GSC”. parvus subzones) at PHE (80 cm) was about one-sixth of that at BBR Class Conodonta Eichenberg, 1930 (4.5 m) (Fig. 30). The thickness of the succeeding asymmetrica- Order Ozarkodinida Dzik, 1976 Norigondolella Subzone of the primitia Zone is ~17 m at BBR, 4.5 Superfamily Gondolelloidea (Lindstrom, 1970) m at PHE, and 32 m at JC, emphasizing the variable paleotopography Family Gondolellidea Lindstrom, 1970 present in the Late Carnian and Early Norian basin. Genus Acuminatella Orchard 2013 At present, correlation with the PM GSSP candidate section in Figures 19, 20; 32-34. Sicily can be achieved at the base of the parvus Subzone, recognized by the nominal species; at a slightly higher level in that subzone where 2013 Acuminatella gen. nov. - Orchard, 2013, p. 446, 447. Parapetella destinae appears; and at the level where typical Carnian Type species: Acuminatella acuminata Orchard. taxa disappear at the top of the parvus Subzone, which correlates with T3 at PM. Additional potential boundary guides recognized in both Diagnosis: The P1 elements of this genus are characteristically arched, regions are Parapetella irwini and Primatella bifida. Other potentially posteriorly downturned, and have a reduced, often pointed posterior useful, short ranging taxa at BBR and PM may be endemics. platform and a prominent carina that extends to the posterior end of the The major decline and extinction of several important Carnian element. Anterior platform margins are weakly to strongly ornate, and conodont genera, the bloom of diminutive taxa, and the ascendency the posterior margins are inornate. The lower surface characteristically of Acuminatella and the Primatella stock, forbearer of Epigondolella, has a pit located in a submedial position and a keel that extends to its is not yet clearly documented at PM. Nevertheless, similar boundary posterior (slightly amended, Orchard, 2013). intervals are recognized. The choice of either Pterosirenites and/or H. Description: The posterior platform gradually tapers or narrows austriaca as Norian indicators, as has been suggested, would have the abruptly at a medial constriction posterior of the anterior nodes. In effect of including some Macrolobatus Zone collections in a newly upper view, tapered forms may have straight, plano-convex, biconvex, defined Norian Stage. or slightly sinuous posterior platform margins. When the platform The sum of data suggests that the base of the asymmetrica- narrowing is abrupt, the posterior platform may be very narrow to Norigondolella Subzone of the primitia Zone offers a position for the vestigial and its margins straight, curved, or sinuous. In all cases, CNB that is most closely aligned with the traditional base of the Norian, the anterior platform margins are raised and each bears 1-6 variably that is, at the base of the Kerri Zone. All Anatropites-bearing collections developed, rounded nodes (in older species) to sharp denticles (in would thereby remain Carnian. At BBR, this level is defined by the younger species). Both the denticles and the platform margins are disappearance of M. parvus and its diminutive associates rather than by covered by compact microreticulae. The free blade is about one-third new appearances, of which there are few; the uncommon Acuminatella element length, highest near the anterior end and forming a low convex curvata may be the only one. The endemic Primatella? gulloae may crest as it descends onto the platform as a row of fairly discrete nodes, fill a similar role at PM. The last M. parvus and its associates close the the largest of which often lie to the posterior. The carina extends to, Carnian chapter in conodont evolution and may be favored as a natural or nearly to, the posterior platform tip. The entire unit is posteriorly boundary. Such a position does, however, remove Halobia species as downarched. A pit is located slightly to the anterior or posterior of definitive indices, and places the range of Pterosirenites on both sides platform midlength, often beneath the constriction if present, and of the CNB. varies in position relative to the keel: it may be subterminal in small ACKNOWLEDGMENTS specimens, but with growth it becomes anteriorly displaced within the pointed keel. This work would not have been possible without the on-going support of the Geological Survey of Canada through a succession of Comparisons: Acuminatella species differ from most contemporaneous projects involving Triassic biostratigraphy, most recently as part of taxa in their reduced posterior platform and strongly developed the Geosciences for Energy and Mapping (GEM) program. Through posterior carina. Some early growth stages of Primatella species many seasons of fieldwork starting in 1981, the late Tim Tozer was (e.g., P. conservativa, P. elongata,) may have reduced posterior a collaborator and mentor on matters Triassic; Dave Gibson also platforms, but these broaden with growth, have a less developed carina provided guidance in the early days, when Frank Riter of Hudson Hope with more discrete nodes, and often a wide posterior platform brim. was the boatman who skilfully provided transport on Williston Lake. Similarly, small specimens of some Quadralella species have reduced More recently, Steve Irwin and Tyler Beatty were able field assistants, posterior platforms, but they also broaden with growth, have less and J-P. Zonneveld was a generous collaborator and logistical lead. developed anterior nodes, and a carina that stops in front of a blunt Leo Krystyn and Chris McRoberts visited the Lake several times and posterior termination. Species of Parapetella with tapered posterior kindly provided expert opinion on, respectively, ammonoid and bivalve platforms have anterior parapets rather than nodes. The Middle Norian faunas. Laboratory and technical support were provided by Peter Orchardella has very high and sharp anterior denticles and less compact Krauss, Hillary Taylor, and Jane Broatch. My family and friends are platform microreticulation. warmly thanked for all their support while this work was brought to Remarks: Kozur (2003) introduced Orchardella for four Middle-Late fruition. Finally, Spencer Lucas, Chris McRoberts, and Bob Nicoll are Norian species formerly referred to Epigondolella that he regarded thanked for their feedback on earlier drafts of the manuscript. as North American endemics (Moix et al., 2007, p. 294); he selected Epigondolella multidentata Mosher, a Middle Norian species, as 42

Figure 32. 1-33. Acuminatella acuminata Orchard 1-3. GSC 132633, sample 18d. 4-6. GSC 132634, sample 21d. 7-9. GSC 132635, sample 18h. 10-12. GSC 132636, sample 21f. 13-15. GSC 132637, sample 18f. 16-18. GSC 132638, sample 21d. 19-21. GSC 132639, sample 22. 22-24. GSC 132640, sample 18. 25-27.GSC 132641, sample 10-03. 28-30. GSC 132615, sample 17b. Holotype. 31-33. GSC 132642, sample PHE-24. Pardonet Hill east. 34-39. Acuminatella curvata sp. nov. 34-36. GSC 132643, sample PHE-24. Pardonet Hill east. 37-39. GSC 132644, sample 23. Holotype. 40-45. Acuminatella binodosa sp. nov. 40-42. GSC 132645, sample PHE-26. Pardonet Hill east. 43-45. GSC 132646, sample 18a. Holotype. Scale bar = 200 microns. 43

Figure 33. 1-3. Acuminatella? prima sp. nov. GSC 132647, sample 1a. Holotype. 4-9. Acuminatella? extensa sp. nov. 4-6. GSC 132648, sample 5. 7-9. GSC 132649, sample 04-21. Holotype. 10-21. Acuminatella sinuosa sp. nov. 10-12. GSC 132650, sample 5. Early form. 13-15. GSC 132651, sample 20f. 16-18. GSC 132652, sample 26. 19-21. GSC 132653, sample 26. Holotype. 22-33. Acuminatella angusta Orchard, alpha morphotype. 22-24. GSC 132654, sample 18h. 25-27. GSC 131166, sample 15. Holotype. 28-30. GSC 132655, sample 18a. 31-33. GSC 131165. sample 14b. 34-41. Acuminatella angusta Orchard, beta morphotype. 34, 35. GSC 132656, sample 14b. 36-38. GSC 132657, sample 17b. 39- 41. GSC 132658, sample 14b. 42-47. Acuminatella longicarinata sp. nov. 42-44. GSC 132659, sample PHE-22. Holotype. Pardonet Hill east. 45-47. GSC 132660, sample 18d. Scale bar = 200 microns. 44

Figure 34. 1-15. Acuminatella constricta sp. nov. 1-3. GSC 132661, sample 17c. 4-6. GSC 132662, sample 17c. 7-9. GSC 132663, sample 17b. 10-12. GSC 132664, sample 13c. Holotype. 13-15. GSC 132665, sample 13a. 16-27. Acuminatella denticulata sp. nov. 16-18. GSC 132666, sample 18d. 19-21. GSC 132667, sample 18e. Holotype. 22-24. GSC 132668, sample 22. 25-27. GSC 132669, sample 18f. 28-48. Acuminatella sagittale sp. nov. 28-30. GSC 132670, sample 18h. 31-33. GSC 132671, sample 17c. 34-36. GSC 132672, sample 9. 37-39. GSC 132673, sample 18c. Holotype. 40-42. GSC 132674, sample 17b. 43-45. GSC 132675, sample 18. 46-48. GSC 132676, sample 8. Scale bar = 200 microns. 45 the type species. A fifth species of Late Carnian-Early Norian age, 2010 Metapolygnathus n. sp. I - Orchard (a), fig. 13. 13-15. originally included within the broad scope of the “Epigondolella 2013 Acuminatella angusta sp. nov. - Orchard, p.448, figs. 4.22-24. primitia population” by Orchard (1983, fig. 2), was regarded by Kozur 2014 Acuminatella angusta Orchard - Balini et al., fig. 1.c1-3.1 (2003, p. 70) as the forerunner of the Middle Norian species but was questionably referred to Orchardella because of the stratigraphic Holotype: GSC 131166, Figure 33. 25-27. gap between the species. The older species is here regarded as a Type stratum: Bed 15, within the Pardonet Formation. homeomorph of Orchardella, and these early forms are referred to Acuminatella (Orchard, 2013). Age: The angusta-dylani Subzone of the primitia Zone. Species of this genus display variation in both platform shape Diagnosis: A species of Acuminatella in which the P1 elements are and anterior node formation but they all have the distinctive narrow narrow and elongate with a straight axis and a denticulate platform posterior platform, and an ornate anterior platform. As in Primatella, mainly developed in the middle one-third of the element. There are evolutionary trends in P1 elements of Acuminatella involve both 2-4 discrete, round, moderately high, and apically pointed nodes on increased differentiation of the anterior platform nodes, and anterior each lateral margin. The anterior one-third consists of a free blade, and migration of the pit. the posterior one-third is a process bearing a very narrow platform or Composition: Eleven species are assigned to Acuminatella, two of lateral flanges. The carina posterior to the indistinct cusp extends to, them with question. The genus is believed to have developed from or nearly to, the posterior end of the element and is composed of 4-6 Quadralella through increased ornamentation of the anterior platform partly to strongly fused and high nodes. The pit underlies the center of margins, prolongation of the carina, and increased lengthening and/or the element, immediately to the posterior of the lateral nodes and far attenuation of the posterior platform. It is hypothesized that advanced anterior of the pointed keel (slightly amended, Orchard, 2013). Quadralella species such as Q. postlobata gave rise to more elongate Comparisons: These elements resemble those of A. sinuosa, but and anteriorly ornate elements that lacked a posterior carina (A.? that species has a distinctly incurved rather than straight posterior prima), and then posterior modification led to the sinuous A.? extensa, platform, a less pronounced posterior carina, and often a relatively the posteriorly reduced A. sinuosa and A. angusta, and a group of longer denticulate anterior platform. Elements of early growth stages elements with tapered platforms starting with A. sagittale. However, of Primatella elongata have a relatively shorter posterior platform that Kraussodontus may provide an alternate origin for the sagittate species. broadens with growth, and a posterior carina that is generally neither as Stratigraphic occurrence: The first, partly questionable representatives high nor as well developed. appear within the spenceri Subzone near the top of the samueli Zone, Remarks: Elements of this species vary in the relative length of the but the genus is largely indicative of the primitia Zone. A succession laterally reduced platform posterior of the marginal nodes, and, on that of species occurs in the Late Carnian through Early Norian, but all basis, two morphotypes are differentiated: alpha morphotype (Fig. 33. disappear immediately prior to the appearance of Epigondolella 22-33) characteristically has 4(-5) posterior carinal denticles, whereas quadrata. The genus is also known in both Haida Gwaii and Nevada. the beta morphotype (Fig. 33. 34-41) has a remarkably long and Acuminatella acuminata Orchard narrow posterior process with 6(-7) denticles. Figure 32. 1-33. Stratigraphic occurrence: The alpha morphotype is abundant in the angusta-dylani Subzone through the upper part of the parvus Subzone. 1983 Epigondolella primitia population - Orchard, fig. 2H, M, N. The beta morphotype is common in the angusta-dylani Subzone 2013 Acuminatella acuminata sp. nov. - Orchard, p. 448, fig. through the acuminata-prominens Subzone of the primitia Zone. 4.16-18. 2013 Acuminatella acuminata Orchard - Carter and Orchard, fig. Acuminatella binodosa sp. nov. 7.13-15. Figure 32. 40-45. 2014 Acuminatella acuminata Orchard - Balini et al., fig. 1.e1,1 Derivation of name: Refers to the single pair of sharp nodes on the f1-2, g1-3. lateral margins of the reduced platform. Holotype: GSC 132615, Figure 32. 28-30. Holotype: GSC 132646, Figure 32. 43-45. Type stratum: Bed 17b, within the Pardonet Formation. Type stratum: Bed 18a, within the Pardonet Formation. Age: The acuminata-prominens Subzone of the primitia Zone. Age: The acuminata-prominens Subzone of the primitia Zone. Diagnosis: A species of Acuminatella in which the relatively long and Diagnosis: These are small P1 elements that have a single sharp node on narrow platform of the P1 element tapers progressively from near its each lateral margin, posterior to which the platform is largely reduced anterior end to the posterior tip; an indentation may occur posterior of or absent. The posterior carina is often composed of two large and high the lateral denticles on one or both margins, particularly in early growth denticles. In profile, the lower surface of the platform is straight to stages. The posterior platform may be somewhat sinuous with one downturned. The pit lies beneath the platform midlength, anterior of margin more convex in outline. Each anterior lateral platform bears 2-3, the narrowly pointed posterior keel. rarely up to five, well developed, discrete, sometimes laterally elongate, and relatively sharp denticles; they may become apically rounded in Comparisons: These P1 elements are homeomorphs of the latest larger specimens. The posterior carina is composed of 2-5 discrete to Triassic Epigondolella bidentata, which differs in having a higher and partly fused, prominent nodes that generally reach the posterior end of sharper anterior denticle pair, a less reduced posterior platform, and a the element, but may terminate a little in front of it. The pit is subcentral more clearly upturned posterior platform. Acuminatella longicarinata in position, commonly lying beneath the anterior half of the platform, sp. nov. has a much longer carina. Primatella sp. nov. A has a lobate and in all cases is anteriorly shifted within the keel (Orchard, 2013). posterior platform. Comparisons: The P1 elements of this species are distinguished from Remarks: Early growth stages of Acuminatella, Primatella, and others referred to Acuminatella by their relative length, the strong Epigondolella often have a “bidentate” morphology and are difficult to denticles of the anterior lateral platform margins, the evenly tapered distinguish. This problem has been discussed in the context of Middle- rather than strongly constricted posterior platform, and the more Late Norian taxa (Orchard, 1983, p. 189), but it is equally applicable anterior position of the pit. A. curvata is broader, shorter, and more around the CNB. The relative size of this species suggests they may uniformly curved on the outer side, although some elements of the be early growth stages of one or more similar species, for example A. present species do approach it (Fig. 32. 31-33). acuminata and A. denticulata. However, elements referred to the former species that are of comparable size already display multiple anterior Stratigraphic occurrence: Abundant from the sagittale-beattyi denticles (e.g., compare with Fig. 32. 1-3). Some A. binodosa occurring Subzone through asymmetrica-Norigondolella sp. Subzone of the with Primatella and Epigondolella may be early growth stages of those primitia Zone. genera. Acuminatella angusta Orchard Stratigraphic occurrence: Very common in the angusta-dylani Figure 33. 22-33; 34-41. Subzone through the basal quadrata Zone 1983[p] Epigondolella primitia population - Orchard, fig. 2O (only). 2007 Metapolygnathus n. sp. I. - Orchard (c), pl. 2, figs. 1-3, 13-15. 46 Acuminatella constricta sp. nov. constricta. Early growth stages resemble those of some Epigondolella Figure 34. 1-15 species, but the platforms of the latter typically broaden and develop Derivation of name: Refers to the strongly constricted posterior much stronger anterior denticles. platform. Stratigraphic occurrence: Common in the lower subdivision of Holotype: GSC 132664, Figure 34. 10-12 parvus Subzone through the asymmetrica-Norigondolella sp. Subzone of the primitia Zone. Type stratum: Bed 13c, within the Pardonet Formation. Acuminatella longicarinata n. sp. Age: The angusta-dylani Subzone of the primitia Zone. Figure 33. 42-47 Diagnosis: A species of Acuminatella in which the P1 element has the 1983 Epigondolella primitia population - Orchard, fig. 2Q. anterior and posterior halves of the platform clearly differentiated by a marked constriction, posterior of which the abruptly narrower posterior Derivation of name: Refers to the long posterior carina. platform tapers to a point. The raised anterior platform margins bear Holotype: GSC 132659, Figure 33. 42-44. poorly developed, irregular, partly coalescing, apically rounded nodes. The pit lies beneath the center of the platform or slightly to its posterior, Type stratum: Bed PHE22, within the Pardonet Formation of east and is subterminal within a rounded loop that may have a short posterior Pardonet Hill. extension of the keel. Age: The asymmetrica-Norigondolella sp. Subzone of the primitia Comparisons: The probable derivative species Acuminatella Zone. denticulata is very similar to this species but differs in having anterior Diagnosis: P1 elements of this Acuminatella species are long and platform ornament composed of sharp denticles rather than blunt nodes. narrow and have a single sharp node on each anterior margin and a This difference is also true of the more posteriorly elongate A. angusta. strongly reduced posterior platform flanking 4-5 large denticles of Unlike the similarly ornate P1 elements of A. sagittale, P1 elements of the posterior carina. In profile, the lower surface of the platform is A. constricta have a strong and persistent medial constriction and the downturned. The pit lies beneath the platform midlength, anterior to the posterior platform is always narrow. narrowly pointed posterior keel. Stratigraphic occurrence: Common in the angusta-dylani Subzone Comparisons: These P1 elements are homeomorphic after the latest through base of middle subdivision of parvus Subzone of the primitia Triassic Epigondolella mosheri, which differs in having a higher and Zone. sharper anterior denticle pair, an often longer carina, and a more clearly Acuminatella curvata sp. nov. upturned posterior platform. They differ from Acuminatella binodosa n. Figure 32. 34-39 sp. in their much longer carina. The carina is equally long in A. angusta, but that possible forbearer has more numerous anterior denticles. Derivation of name: Refers to the curved axis of the P1 element. Stratigraphic occurrence: Common in the angusta-dylani Subzone Holotype: GSC 132644, Figure 32. 37-39 through asymmetrica-Norigondolella sp. Subzone of the primitia Zone. Type stratum: Bed 23, within the Pardonet Formation Acuminatella sagittale sp. nov. Age: Low in the asymmetrica-Norigondolella sp. Subzone of the Figure 34. 28-48 primitia Zone. Derivation of name: From sagitt (Latin), referring to the arrow- Diagnosis: P1 elements of this Acuminatella species have a distinctly shaped platform. curved carina and a relatively broad platform with a plano-convex to Holotype: GSC 132673, Figure 34. 37-39 biconvex outline. The anterior marginal nodes are discrete but may be laterally elongate and of variable height and sharpness. The low Type stratum: Bed 18c, within the Pardonet Formation. nodes of the carina, which may be partly fused, extend to the posterior Age: Lower subdivision of parvus Subzone of the primitia Zone. platform tip. The pit lies beneath the center of the platform or a little posterior of that point, and the keel extends posteriorly beyond it. Diagnosis: A species of Acuminatella in which the lateral platform margins of the evenly tapered P1 elements have a generally weakly Comparisons: These P1 elements are shorter and broader than in biconvex or plano-convex outline, although some smaller specimens comparably sized elements of A. acuminata (q.v.), which have a straight may have a medial platform indentation. The raised anterior lateral to weakly curved axis. margins bear poorly to moderately well developed, commonly discrete, Stratigraphic occurrence: Rare and only known from the asymmetrica- apically rounded nodes. The posterior platform tapers rapidly in its Norigondolella sp. Subzone of the primitia Zone. posterior ¼ to a narrowly rounded to pointed tip. The carina reaches to, or close to, the posterior end of the platform from which it may Acuminatella denticulata sp. nov. be separated by a very narrow brim. The pit is submedial in position Figure 34. 16-27 with respect to the platform, and with growth it becomes increasingly Derivation of name: Refers to the strong denticles of the anterior displaced anteriorly with respect to the keel. platform. Comparisons: The posterior platform is much broader, more evenly Holotype: GSC 132667, Figure 34. 19-21. tapered, and lacks the dual medial constriction of Acuminatella constricta. The platform of the younger A. acuminata is relatively Type stratum: Bed 18e, within the Pardonet Formation. longer, the anterior marginal denticles much more defined and apically Age: Lower subdivision of parvus Subzone of the primitia Zone. sharp, and the pit commonly more anterior in position. Diagnosis: A species of Acuminatella with P1 platforms that are Stratigraphic occurrence: Abundant in the sagittale-beattyi Subzone differentiated by a medial constriction into an anterior, sharply through middle subdivision of the parvus Subzone of the primitia Zone. denticulate part and a posterior part that is narrow and unornamented. Acuminatella sinuosa sp. nov. Small specimens have a short, narrowly triangular platform with 1 or 2 Figure 33. 10-21 anterior marginal denticles, whereas in larger specimens a characteristic constriction occurs posterior to 2-4 marginal denticles. The posterior Derivation of name: Refers to the distinctly curved posterior carina, carina commonly has a large penultimate node near the center of the which imparts a sinuous appearance to the platform. posterior platform followed by a smaller, or incipient node near the Holotype: GSC 132653, Figure 33. 19-21. pointed posterior end. The pit lies beneath the center of the platform in mature growth stages, anteriorly displaced within the keel. Type stratum: Bed 26, within the Pardonet Formation. Comparisons: These elements are relatively shorter than those of the Age: The asymmetrica-Norigondolella sp. Subzone of the primitia similarly denticulate Acuminatella acuminata, adults of which differ Zone. also in lacking a strongly constricted posterior platform. The sharp Diagnosis: P1 elements of this Acuminatella species have an anterior anterior denticles contrast with the low nodes of the similarly shaped A. and posterior platform of subequal length and a distinctly curved 47 posterior part. There are 2-4 well developed, discrete and generally fewer, larger, and generally less discrete anterior marginal platform sharp denticles on each anterior lateral margin, posterior to which there nodes. Unlike typical Acuminatella species, the carina does not extend is a marked narrowing of the platform. The curved posterior process is to the posterior margin. Unlike A.? extensa (q.v.), the pit is posteriorly ½ as wide as the anterior platform and bears 3-4 larger carinal denticles located and is terminal in the keel. that extend close to the posterior tip. The pit lies beneath the platform Remarks: This rare species seems to occupy an intermediate position midlength opposite the constriction, and within a keel that extends a both morphologically and stratigraphically between Quadralella variable distance to its posterior. species with their posterior brim, and Acuminatella with their more Comparisons: The curved posterior process of the P1 element of this developed anterior nodes. species serves to distinguish it from Acuminatella angusta, which also has a more reduced anterior platform and a longer posterior part. Stratigraphic occurrence: Rare in the medioconstricta Subzone of the Elements of A.? extensa have a broader and more asymmetrically samueli Zone. developed posterior platform and lack a long posterior carina. Genus Carnepigondolella Kozur 2003 Remarks: The first and less common examples of this species are from Figures 17, 18; 35-39 low in the succession (e.g., Fig. 33. 10-12) and have more apically 2003 Carnepigondolella - Kozur, p. 68-69. rounded anterior platform denticles compared with those from younger 2012 Carnepigondolella Kozur - Mazza et al. (b), p. 92-3. stratigraphic levels. 2013 Carnepigondolella Kozur - Orchard, p. 448-9. Stratigraphic occurrence: Common in the sagittale-beattyi Subzone Type species: Metapolygnathus zoae Orchard 1991b. through asymmetrica-Norigondolella sp. Subzone of the primitia Zone. Description: In upper view, the P1 platform is commonly subrectangular Acuminatella? extensa sp. nov. with a squared-off posterior margin that is less commonly symmetrically Figure 33. 4-9 or asymmetrically expanded postero-laterally, or posteriorly rounded. Derivation of name: Refers to the extended posterior platform of the In some species, a platform constriction commonly occurs in a posterior P1 element. (stratigraphically older species) to medial (younger species) position. In profile, the lower edge of the element is downcurved in the posterior Holotype: GSC 132649, Figure 33. 7-9. third but may appear straight in the youngest species when viewed from Type stratum: Bed 04-21 (approximates beds 5-6), within the Pardonet the inner side. Formation. The relatively flat platforms bear low distinct rounded nodes and/ or sharp denticles on the anterior platform margins and often also on Age: The sagittale-beattyi Subzone of the primitia Zone. the posterior margins. When viewed in profile, the nodes or denticles Diagnosis: The distinctive P1 element has a long, broad, strongly are usually between 10% and 30% of the total depth of the element inturned and asymmetric posterior platform. The anterior ½ of the measured from the node/denticle tip to the basal keel margin. A dense platform bears 3-6 moderately discrete but closely spaced nodes that microreticulation occurs on all platform margins and nodes and is only are apically more rounded and less differentiated to the posterior. The absent from sharp denticles. carina is composed of 6 low and discrete nodes that terminate in a large Stratigraphically older species have short blades about one-quarter node immediately in front of a long, inwardly deflected tongue. The element length; the youngest have longer blades equal to one-half lower surface has a pit located at the junction of the nodose anterior element length. The carina generally terminates well in front of the end part and posterior platform, beyond which there is a long, sinuous, and of the platform, and a wide posterior brim is developed. A distinctive pointed keel. cusp is often the last node of the carina, but it may be subterminal and Comparisons: The strong posterior asymmetry of these nodose indistinct. The pit is situated beneath the posterior part of the platform elements sets them apart from most other contemporaneous elements. in stratigraphically older species and beneath the platform midlength in Less pronounced posterior asymmetry is displayed by Acuminatella younger species; it lies anterior to the rounded to bifid keel termination. sinuosa, which has a much narrower posterior platform and a carina Comparisons: Compared with the P1 elements of both Primatella and that extends to the posterior margin. The presence of a posterior Epigondolella, those of Carnepigondolella are generally smaller and platform brim, a feature of both A.? extensa and the similar A.? prima, have less robust and often more numerous platform nodes/denticles. is typical of Quadralella species, but those species rarely have the well- The platform margins anterior to the denticles in Primatella are developed anterior nodes typical of Acuminatella. usually much steeper, and the denticles/nodes tend to be well formed Remarks: The present species, and Acuminatella? prima, are assigned and of comparable size rather than gradually increasing in size and with question to the present genus because they retain a posterior height towards the posterior as in Carnepigondolella. The denticles platform brim, reminiscent of ancestral Quadralella species. The of Epigondolella species are even larger, and the lower profiles of two species appear to represent transitional forms in which posterior those elements are commonly stepped upward posteriorly. Typical platform modification resulted in progressive elongation of the posterior Acuminatella species differ in having pointed platforms and a well- platform. developed posterior carina. Platform microreticulation patterns are similar in Carnepigondolella, Acuminatella, and Primatella, but differ Stratigraphic occurrence: Rare in the sagittale-beattyi Subzone, in Epigondolella (q.v.). ?angusta-dylani Subzone of the primitia Zone. Remarks: Orchard (1991a) described a distinctive group of Acuminatella? prima sp. nov. Upper Carnian ornate elements from Haida Gwaii as species of Figure 33. 1-3 Metapolygnathus: M. zoae, M. samueli, and M. pseudoechinatus. Derivation of name: From prim- (Latin), meaning first, alluding to the Later, Kozur (2003, p. 68-69) assigned these to a new genus, species representing the initial step in the development of Acuminatella. Carnepigondolella. Contrary to Kozur’s classification, the species lindae and nodosus are here excluded from Carnepigondolella (see Holotype: GSC 132647, Figure 33. 1-3. Quadralella). This is true also for the elements previously assigned to Type stratum: Bed 1a, within the Pardonet Formation. the species Metapolygnathus pseudoechinatus (see Primatella); those elements are now assigned to C. medioconstricta and C. spenceri. Age: The medioconstricta Subzone of the samueli Zone. At Black Bear Ridge, common representatives of Diagnosis: The P1 element has a relatively long platform in which the Carnepigondolella occur in the Ludington Formation and basal Pardonet Formation, where a succession of elements displays a clear posterior 1/3 of the platform abruptly narrows to produce a terminally rounded lobe, at the center of which there is a large terminal cusp morphological trend in the reduction of platform length, concomitant surrounded by a platform brim. Anterior of the lobe, the lateral platform lengthening of the free blade, and anterior migration of the pit (Orchard, margins bear 4-6 small (~¼ of platform height), closely spaced, and 2007c, p. 132-3). Some species also have anterior denticles that apically rounded nodes. The pit is located beneath the cusp at the transition into posterior nodes, and a further trend appears to be toward posterior end of a grooved keel. increasing the former at the expense of the latter. Several lineages are postulated here (Figs. 17, 18). That including the type species, C. zoae, Comparisons: This species has a longer and narrower anterior platform may be the only one eventually retained in Carnepigondolella. than its probable precursor, Quadralella postlobata, which also has Two exceptions to the previous scope of the genus are noted. Mazza 48 et al. (2010, 2012b) noted that the pit in Carnepigondolella always lay Age: The eozoae-ludingtonensis Subzone of the samueli Zone. “distinctly behind” the platform midpoint, whereas advanced species Diagnosis: The P1 element of this Carnepigondolella species has well here assigned to the genus do have a pit that lies at, or even anterior to defined marginal platform nodes that are well rounded in both upper the platform midpoint (e.g., C. spenceri). A second exception may be and lateral views, and which are more developed in the anterior half represented by a single contemporary element (C.? sp. nov. A), which of the elongate platform, becoming less differentiated towards the displays typical Carnepigondolella anterior denticles but, atypically, posterior. A constriction occurs in the posterior 1/5 of the platform. The both a posterior carina and a pointed platform and keel (contra Kozur, free blade is about ⅓ of the total element length, forms a convex crest, 2003, p. 68). The youngest representatives of Carnepigondolella at Black Bear and passes into a low carina that extends to the centre of the posterior Ridge are relatively diminutive elements that appear at about the same constricted part of the platform; the cusp is indistinct. The pit lies at the time as the appearance of quite different species of Acuminatella (Figs. posterior end of both the keel and the downturned platform. 19, 20) and Primatella (Fig. 22); these latter are believed to have arisen Comparisons: The platform nodes of the new species are identical to from Quadralella (q.v.). Both Kozur (2003) and Mazza et al. (2010, p. those of Carnepigondolella zoae but the latter has a shorter platform, a 123; 2012a, p. 19; 2012b, p. 92) regarded Carnepigondolella as ancestral platform constriction in a submedial position, and, in the holotype, an to Epigondolella, but the revised concepts of these two genera separate anteriorly shifted pit that lies slightly posterior of platform midlength. them stratigraphically, and the latter is hypothesized to have arisen The far posterior position of the constriction in C. eozoae is unlike any directly from Primatella and to be unrelated to Carnepigondolella. other species of the genus. Composition: Twelve species of Carnepigondolella and eight Remarks: Carnepigondolella zoae is envisaged to have arisen from C. variations or morphotypes are differentiated from Black Bear Ridge. eozoae through anterior shifting of both the platform constriction and Mazza and Rigo (in Mazza et al., 2012b) referred several additional the pit. A specimen tentatively included in C. eozoae, from east Pardonet species to Carnepigondolella but they are excluded from that genus, Hill (Fig. 35. 1-3), has a reduced platform and a marked posterior and assigned to Quadralella (e.g., Q. carpathica) or Primatella (e.g., constriction. This element may represent a juvenile morphology from P. orchardi). which, it is envisaged, later growth produced lengthening of the anterior Stratigraphic occurrence: The present concept of this genus restricts platform. A possible ancestor of the present species, here called C. aff. typical species of Carnepigondolella to the Upper Carnian samueli eozoae (Fig. 17), occurs at a lower stratigraphic level in Haida Gwaii: Zone, and undetermined older strata. The holotypes of both C. zoae and it has a very similar platform shape but less developed anterior nodes. C. samueli came from Haida Gwaii where they were bracketed between Stratigraphic occurrence: Common in the eozoae-ludingtonensis ammonoids of the upper Welleri Subzone and the Macrolobatus Zone Subzone to zoae Subzone of the samueli Zone. (Orchard 1991a, fig. 5; 1991b, p. 318, 319). Carnepigondolella gibsoni sp. nov. Carnepigondolella anitae sp. nov. Figure 36. 1-8, ?9-11 Figure 35. 16-23 Derivation of name: Named for David Gibson in tribute to his regional 1991 Metapolygnathus n. sp. G - Orchard (a), p. 176, 189, pl. 1, fig. Triassic studies in northeastern British Columbia. 18. 2010 Carnepigondolella zoae (Orchard) - Mazza et al., pl. 1, figs. Holotype: GSC 132685, Figure 36. 6-8. 7a-c. Type stratum: Bed H, within the Ludington Formation. 2010 Carnepigondolella zoae B - Mazza et al., pl. 1, figs. 8a-c. Age: The eozoae-ludingtonensis Subzone of the samueli Zone. Derivation of name: Named for Anita Lecesse, in gratitude for her support during the course of this study. Diagnosis: These Carnepigondolella P1 elements are relatively narrow and long, with a length-to-breadth ratio of about 3:1. A Holotype: GSC 132682, Figure 35. 21-23. nodose or denticulate anterior platform is about twice the length of Type stratum: Bed 0a, within the Ludington Formation. the unornamented posterior part, which is separated by a constriction in early growth stages; later growth stages have subparallel margins. Age: The zoae Subzone of the samueli Zone. The 3-5 anterior platform nodes are sharp but may pass into lower, Diagnosis: The P1 element is relatively slender and bears small sharp more rounded nodes to the posterior. The free blade is short, ¼ of total denticles on the margins of the anterior ⅓ of the platform, rounded element length, and passes into a row of 4-6 carinal nodes that extend to nodes on the medial ⅓ of the platform, and no ornament on the the middle of the posterior platform without displaying an exceptional relatively narrow, constricted posterior ⅓ of the platform. A large cusp cusp. The pit lies near the posterior end of the platform and keel. is situated on the posterior platform and may have a second smaller Comparisons: This species resembles Carnepigondolella samueli in its node to its posterior separated from the posterior margin by a narrow relatively short blade and posterior pit position, but it is relatively longer platform brim. The pit lies at the posterior end of the platform and keel. and lacks the strong posterior nodes. It differs from C. pseudodiebeli, Comparisons: This species displays platform ornament that is a which also has an unornamented posterior platform, in its longer and combination of that seen in C. zoae, characterized entirely by rounded narrower platform. Although small specimens may show a platform nodes, and that typical of C. samueli, characterized by wholly sharp constriction, the species lacks the stronger medial constriction, denticles. posterior expansion, and longer blade typical of C. medioconstricta and the strong posterior constriction of C. eozoae. However, one element Remarks: This species appears to demonstrate one evolutionary trend (Fig. 36. 9-11) does have more rounded nodes and a slight posterior in some Carnepigondolella that involves the progressive development constriction and as such appears transitional to C. eozoae. of sharp platform denticles starting from the anterior platform margins and proceeding towards the posterior. Remarks: These elements co-occur with Carnepigondolella samueli The through much of the exposed Ludington Formation at Black Bear Ridge specimens illustrated by Orchard (1991a) and Mazza et al. but do not range as high. Broader elements resembling C. samueli in (2010) exhibit the intermediate features of this new species. shape but lacking posterior ornament like the present species appear Stratigraphic occurrence: Uncommon in the zoae Subzone to later and are assigned to P. pseudodiebeli. medioconstricta Subzone of the samueli Zone. The specimens from Pizzo Mondello (Mazza et al., 2010, samples PM19, FNP53A) occur Stratigraphic occurrence: Common from the base of section through within a comparable interval. the zoae Subzone of the samueli Zone. Carnepigondolella eozoae Orchard Carnepigondolella medioconstricta Orchard Figure 35. 1-3?, 4-6 Figure 37. 1-23 Carnepigondolella medioconstricta sp. nov 2013 Carnepigondolella eozoae sp. nov. - Orchard, 2013, p. 449, 2013 . - Orchard, p. figs. 3.1-3.3. 449, figs. 3.10-3.12. 2013 Carnepigondolella medioconstricta Orchard - Carter and Holotype: GSC 132598, Figure 35. 4-6. Orchard, figs. 3. 7-9. Type stratum: Bed C, within the Ludington Formation. Holotype: GSC 132601, Figure 37. 16-18. 49

Figure 35. 1-3?, 4-6. Carnepigondolella eozoae Orchard. 1-3? GSC 132677, sample PHE-13a. Pardonet Hill east. 4-6. GSC 132598, sample C. Holotype. 7-15. Carnepigondolella zoae (Orchard). 7-9. GSC 132678, sample 0a. 10-12. GSC 132679, sample 04-108A. 13-15. GSC 95203. Holotype. GSC cur. no. C-157037, Peril Formation. Huston Inlet. Haida Gwaii. 16-23. Carnepigondolella anitae sp. nov. 16, 17. GSC 132680, sample 0. 18-20. GSC 132681, sample 1a. 21-23. GSC 132682, sample 0a. Holotype. Scale bar = 200 microns.

Type stratum: Bed 2, within the Pardonet Formation. denticles that commonly terminate in a large cusp at the centre of the Age: The medioconstricta Subzone of the samueli Zone. posterior platform; a smaller carinal node may occur to its posterior within an otherwise broad platform brim. The pit lies at or posterior of Diagnosis: Carnepigondolella P1 elements have a relatively elongate, the centre of the platform and is anteriorly displaced within the keel. somewhat curved platform with a marked medial constriction, particularly in smaller growth stages, and a posterior platform that is Comparisons: The rounded posterior outline differs from that seen in laterally expanded, particularly on the inner side. The anterior platform contemporaneous species other than in Carnepigondolella spenceri, generally bears 2-4 discrete and sharp nodes on each lateral margin, which has a much shorter and often posteriorly expanded platform. C. whereas the often asymmetrical, triangular to semicircular posterior medioconstricta may be somewhat rounded but its slender constricted platform is unornamented or has weakly developed marginal nodes. outline sets it apart. Some elements of the younger species of Primatella The free blade is long, equal to one-half the element length in small rotunda resemble C. milanae, but the former can be distinguished by its specimens and relatively a little shorter in larger specimens. One or larger and more irregularly developed anterior nodes formed on more two carinal nodes occur on the posterior platform, anterior to a wide upturned platform margins. platform brim. The cusp is inconspicuous above a pit that underlies the Remarks: See C. aff. milanae. center of the platform; the posterior keel may be bifurcated (Orchard, 2013). Stratigraphic occurrence: Very common in the medioconstricta Subzone through spenceri Subzone of the samueli Zone. Comparisons: The present species resembles Primatella orchardi, but that younger species is often larger, has more robust anterior platform Carnepigondolella aff. milanae nodes, a relatively shorter blade, more discrete carinal nodes, and it Figure 38. 12-14 lacks incipient nodes on the posterior margins. Remarks: This unique specimen differs from typical Carnepigondolella Stratigraphic occurrence: Abundant in the medioconstricta Subzone milanae of comparable size in the medial position of its pit and long through spenceri Subzone of the samueli Zone. The species occurs also posterior keel, the relatively short carina, and the laterally broadened in a similar association on Haida Gwaii (Carter and Orchard, 2013). anterior denticles. These differences in pit position are comparable to that which distinguishes the morphotypes of Primatella rotunda and, Carnepigondolella milanae sp. nov. like that species, the more anteriorly shifted pit is seen near the upper Figure 38. 1-11 limit of its stratigraphic range. Derivation of name: Named for Milana, my grand-daughter. Stratigraphic occurrence: Rare and only known from medioconstricta Holotype: GSC 132706, Figure 38. 9-11 Subzone of the samueli Zone. Type stratum: Bed 4, within the Pardonet Formation. Carnepigondolella postsamueli sp. nov. Figure 39. 24-28; 29, 30 Age: The spenceri Subzone of the samueli Zone. 2007 Carnepigondolella n. sp. N - Orchard (b), figs. 1.13-15. Diagnosis: A species of Carnepigondolella in which the P1 element has an elongate-oval platform with a distinctive unexpanded, rounded Derivation of name: Referring to its development from C. samueli. posterior margin lacking ornamentation, and anterior platform margins Holotype: GSC 132724, Figure 39. 26-28. that bear sharp to rounded nodes. The free blade is equal to about ⅓ to ½ total element length and continues onto the platform as a row of low Type stratum: Bed 1c, within the Pardonet Formation. 50

Figure 36. 1-8, ?9-11. Carnepigondolella gibsoni sp. nov. 1, 2. GSC 132683, sample H. 3-5. GSC 132684, sample 0. 6-8. GSC 132685, sample H. Holotype. ?9-11. Element transitional to C. eozoae. GSC 132686, sample G. 12-21. Carnepigondolella pseudodiebeli (Kozur), alpha morphotype. 12, 13. GSC 132687, sample 2a. 14, 15. GSC 132688, sample 1. 16-18. GSC 132600, sample E. 19, 20. GSC 132689, sample 2a. 21. GSC 132690, sample G. 22-31. Carnepigondolella pseudodiebeli (Kozur), beta morphotype. 22, 23. GSC 132691, sample E. 24-26. GSC 132692, sample E. 27, 28. GSC 132693, sample E. 29-31. GSC 132694, sample 0. Scale bar = 200 microns.

Age: The medioconstricta Subzone of the samueli Zone. Stratigraphic occurrence: Abundant in the medioconstricta Subzone through spenceri Subzone of the samueli Zone. Diagnosis: The P1 elements have a subrectangular, symmetrical to asymmetrical platform with a length-to-breadth ratio of between 2:1 Carnepigondolella pseudodiebeli (Kozur) and 2.5:1, and a free blade that is between ⅓ and ½ element length. Figure 36. 12-21; 22-31 Marginal denticles occur throughout the platform, are short and sharp 1972 Metapolygnathus spatulatus pseudodiebeli n. subsp. - Kozur, in the anterior half, and more obliquely oriented in the posterior part, p. 8-9, pl. 4, figs. 5a, b. especially on the posterior margin. The pit lies a little posterior of 2010 Carnepigondolella pseudodiebeli (Kozur) - Mazza et al., pl. midlength and in front of the posterior end of the often bifurcated keel. 1, figs. 9a-c. Comparisons: Carnepigondolella samueli and the present species can 2012[p] Carnepigondolella pseudodiebeli (Kozur) central be distinguished by the relatively greater platform length and shorter morphotype - Mazza et al. (b), pl. 2, figs. 6a-c, 10a-c (only). free blade of the former, although specimens with broken blades may Description: A species of Carnepigondolella in which P1 elements only be distinguishable by the more forward shifted pit in the present have a relatively short, subrectangular platform with a length-to- species. Large asymmetrical specimens of C. medioconstricta may breadth ratio of between 1.5:1 and 2:1, well differentiated and often have a similar shape to the beta morphotype (see below), but those laterally broadened denticles on each anterior platform margin, and a elements bear no ornament on the posterior platform margin, and variable but often very short free blade. Small specimens may exhibit a typical specimens have a strong medial platform constriction. platform constriction in the posterior ⅓ of the platforms beyond which Remarks: Two morphotypes are distinguished that parallel those they are laterally expanded; this expansion can be very pronounced at of Carnepigondolella samueli, namely an alpha morphotype (Fig. the posterior end of large specimens. The carina commonly terminates 39. 24-28) with a symmetrical platform, and the relatively rare beta in a variably conspicuous cusp at the beginning of the posterior morphotype (Fig. 39. 29, 30) with an asymmetrical platform. expansion, and secondary carina may be formed in large elements. The pit lies close to the posterior end of the platform and of the keel, which 51

Figure 37. 1-23. Carnepigondolella medioconstricta Orchard. 1, 2. GSC 132695, sample 1a. 3, 4. GSC 132696, sample 2. 5-7. GSC 132697, sample 2. 8, 9. GSC 132698, sample 2. 10-12. GSC 132699, sample 2. 13-15. GSC 132700, sample 1c. 16-18. GSC 132601, sample 2. Holotype. 19, 20. GSC 132701, sample 1a. 21-23. GSC 132702, sample 4. Scale bar = 200 microns. is typically bifurcated in moderate to large specimens. the lower margin is usually more upturned posteriorly. The elements assigned to Carnepigondolella orchardi by Mazza Remarks: This species has been broadly interpreted by previous et al. (2012b) are shorter than the holotype of that species and are more authors, and the present concept also embraces some variation that is posteriorly expanded, as in the present species. The recently described difficult to evaluate in the absence of a well described and illustrated Epigondolella heinzi Mazza, Cau and Rigo, also appears more typical type population. Mazza et al. (2012b) emphasized the bifid keel as a of a Carnepigondolella; it differs in being more uniformly expanded diagnostic feature of Carnepigondolella pseudodiebeli and consequently posteriorly and has a more medial pit. included some elongate elements with more subdued anterior nodes (their morphotype A) in the species. However, these elements are here Stratigraphic occurrence: Uncommon in the eozoae-ludingtonensis referred to Quadralella kathleenae (q.v.). Bifid keels are commonly Subzone through zoae Subzone of the samueli Zone. Mazza et al. developed in large quadrate elements of several different genera and (2012b) record their elements from the Tuvalian. species and are not regarded as grounds for taxonomic differentiation Carnepigondolella samueli (Orchard 1991) (e.g., Ancyrogondolella Budurov 1972). Two morphotypes of C. Figure 39. 1-12; 13-20 pseudodiebeli are differentiated at Black Bear Ridge: 1989 “Epigondolella” n. sp. G - Orchard, in Carter et al., pl. 1, fig. alpha morphotype 6. Figure 36. 12-21 1991 Metapolygnathus n. sp. G - Orchard (a), p. 176, pl. 1, figs. Remarks: These are relatively long elements with transversely 12-18; pl. 3, figs. C, D. elongated anterior nodes and are close to the type material. The P1 1991 Metapolygnathus samueli n. sp. - Orchard (b), p. 318-19, pl. elements of Carnepigondolella samueli and C. postsamueli are similar 1, figs. 10-12. but have well developed posterior platform nodes, whereas those of C. 2007 Carnepigondolella samueli (Orchard) - Orchard (b), fig. gibsoni have longer and narrower platforms. 1.22-24. 2012 Carnepigondolella samueli (Orchard) - Mazza et al. (b), p. Stratigraphic occurrence: Abundant in the eozoae-ludingtonensis 98-100, pl. 3, fig. 1, 2. Subzone through spenceri Subzone of the samueli Zone. 2012 [?] Epigondolella vialovi (Buryi) - Mazza et al. (b), p. 111, pl. 7, beta morphotype figs. 2, 3. Figure 36. 22-31 Holotype: GSC 81244, Figure 39. 7-9. 2007 Carnepigondolella n. sp. E - Orchard (c), p. 136. Description: A species of Carnepigondolella in which the P1 platform 2010 [?] Carnepigondolella orchardi (Kozur) - Mazza et al., pl. 1, element is characterized by a short free blade equal to about ¼ total figs. 11a-c. element length, and by small, sharply terminated nodes on all the 2012 [?] Carnepigondolella orchardi (Kozur) - Mazza et al.(b), p. 96- platform margins. A slight constriction is common in the posterior one- 7, pl. 2, figs. 1a-c, 2a-c. third of the platform, beyond which the marginal nodes may be less Remarks: These elements are shorter than the alpha morphotype and well developed. The posterior platform is typically rectangular but may have higher, less laterally expanded nodes. These elements resemble be postero-laterally expanded, particularly in late growth stages; this the younger Primatella orchardi and P. subquadrata, but the blades of expansion may be asymmetric. In profile, the blade forms a low convex those species are relatively longer, and the pit lies in a more anterior crest that extends onto the platform as a row of low carinal nodes that position. Also, incipient nodes may appear on the posterior margin of generally terminate in the centre of the expanded posterior platform, the present specimens, and the anterior nodes are generally smaller than well in front of the posterior margin. The pit is located close to the in Primatella, although they may be more discrete and apically sharper. posterior platform margin, close to the posterior end of the prominent The anterior denticles of Epigondolella quadrata are much longer, and 52

Figure 38. 1-11. Carnepigondolella milanae sp. nov. 1, 2. GSC 132703, sample 2. 3-5. GSC 132704, sample 2a. 6-8. GSC 132705, sample 1c. 9-11. GSC 132706, sample 4. Holotype. 12-14. Carnepigondolella aff. milanae. GSC 132707, sample 2a. 15-19. Carnepigondolella aff. spenceri. 15-17. GSC 132708, sample 1a. 18, 19. GSC 132709, sample 4. 20-35. Carnepigondolella spenceri sp. nov. 20-22. GSC 132710, sample 4. 23. GSC 132711, sample 4. 24-26. GSC 132712, sample 4. Holotype. 27, 28. GSC 132713, sample 3. 29-31. GSC 132714, sample 4. 32. GSC 132715, sample 4. 33-35. GSC 132716, sample 3. Scale bar = 200 microns. 53

Figure 39. 1-12. Carnepigondolella samueli (Orchard), alpha morphotype. 1-3. GSC 132717, sample P. 4-6. GSC 132599, sample 0a. 7-9. GSC 81244. Holotype. GSC cur. no. C-157037, Peril Formation. Huston Inlet. Haida Gwaii. 10-12. GSC 132718, sample C. 13-20. Carnepigondolella samueli (Orchard), beta morphotype. 13-15. GSC 132719, sample 4. 16-18. GSC 132720, sample 0. 19, 20. GSC 132721, sample E. 21-23. Carnepigondolella? sp. nov. A. GSC 132722, sample 0. 24-28. Carnepigondolella postsamueli sp. nov., alpha morphotype. 24, 25. GSC 132723, sample 2. 26-28. GSC 132724, sample 1c. Holotype. 29, 30. Carnepigondolella postsamueli sp. nov., beta morphotype. GSC 132725, sample 4. 31-34. Carnepigondolella aff. spatulata (Hayashi). 31, 32. GSC 132726, sample 4. 33, 34. GSC 132727, sample 4. Scale bar = 200 microns. keel, which may be bifurcated in large specimens. belong here. Comparisons: The species is very close to Carnepigondolella Two morphotypes are distingueshed in this work: an alpha postsamueli but differs in having a shorter blade and more posteriorly morphotype (Fig. 39. 1-12) with a rectangular platform, and a beta located pit. It differs from C. pseudodiebeli in its well-developed morphotype (Fig. 39. 13-20) with a generally larger, asymmetrical posterior ornament. platform in which the inner postero-lateral margin is laterally expanded. Remarks: The holotype of this species (Fig. 39. 7-9), from the Stratigraphic occurrence: This species is the index to the Upper Peril Formation of Huston Inlet, Haida Gwaii, has strong denticles Carnian samueli Zone. At its type locality, the species is not surrounding the entire perimeter of the platform and identical elements independently dated, but nearby on Kunghit Island, the species occurs occur at Black Bear Ridge. Larger specimens of Carnepigondolella within the Late Carnian above ammonoids of the Welleri Subzone II samueli with more subdued posterior ornament are close to the large and below those assigned to the Macrolobatus Zone. At Black Bear holotype of C. pseudodiebeli Kozur, but both species are retained based Ridge, abundant alpha morphotype ranges from the base of the section on this difference. Mazza et al. (2012b) identified this species in Sicily, (pre- eozoae-ludingtonensis Subzone) through the spenceri Subzone, but noted that the Neotethyan examples were less slender and had whereas the uncommon beta morphotype was recovered from the more numerous posterior nodes; they also differentiated a Morphotype eozoae-ludingtonensis Subzone through the zoae Subzone. In Sicily, C. A, which differs most clearly in its more rounded posterior platform. samueli ranges in the Upper Tuvalian (Mazza et al., 2012b). One specimen identified asE. vialovi by Mazza et al. (2012b) may also Carnepigondolella aff. spatulata (Hayashi) 54 Figure 39. 31-34 report, also differ in the presence of a strongly ornate posterior platform 1968 [?] Gladigondolella abneptis var. spatulata var. nov. - Hayashi margin. Compared with the younger and equally squat Primatella ex (a), p. 69, pl. 2, fig.5. gr. pseudoechinata, the P1 element of C. spenceri broadens posteriorly 1991 Metapolygnathus echinatus (Hayashi) - Orchard (a), p. 189, rather than narrows, and it has smaller anterior nodes. Primatella pl. 1, fig. 25. subquadrata, which also has a broad posterior platform brim, has a 2010 [?] Epigondolella spatulata (Hayashi) - Mazza et al., pl. III, figs. longer platform, shorter blade, and larger anterior nodes. Epigondolella 5a-c, ?6a-c. quadrata has a more angular posterior outline and much larger anterior nodes. Description: The P1 elements have a relatively short equidimentional platform with a length-to-breadth ratio of ~1:1, strong nodes on both Remarks: This species and allied forms appear to be the end members lateral and posterior margins, and a free blade that is equal to or a of the Carnepigondolella lineage with its trend of anterior platform little longer than the platform. The platform is generally rectangular in reduction and the resulting anterior shift of the pit. The Sicilian species outline, but the smallest specimen has a subrounded posterior margin. spatulata, cf. spatulata, and heinzi, which were assigned to the genus The pit underlies the center or anterior part of the platform within a Epigondolella on the basis of their central pit and shorter platform, all broad keel that expands to the posterior. have relatively small and low denticles, unlike true Epigondolella (q.v.) but resembling more this group of reduced Carnepigondolella species. Comparisons: These elements closely resemble the holotype of Epigondolella cf. spatulata sensu Mazza et al. (2012b) appears very Epigondolella spatulata but differ in their more rectangular rather than close to C. spenceri but differs in its subcircular platform outline; also, rounded posterior outline, and with a maximum width at the posterior it apparently comes from a much higher, upper Lacian level. end rather than in the middle of the platform. The similarly squat Carnepigondolella balogni (sensu Mazza et al., 2012b, pl. 1, fig. 1) also Stratigraphic occurrence: Abundant in the spenceri Subzone of the differs in its rounded platform as well as a shorter free blade, expanded samueli Zone. anterior trough margins, and wide anterior nodes. Carnepigondolella aff. spenceri Remarks: Carnepigondolella postsamueli is probably the direct Figure 38. 15-19 ancestor of this species, which differs in its longer blade and shorter Remarks: These elements are less common than Carnepigondolella platform. A possibly transitional form with a reduced platform is spenceri and appear earlier. They differ in being relatively longer with included here (Fig. 39. 31, 32). The shortest platforms occur in elements a subtriangular shaped posterior platform, relatively angular postero- from the youngest stratigraphic levels. lateral corners, and occasional incipient nodes on the posterior margin. Epigondolella spatulata, although broadly interpreted in the past, has been more narrowly defined by Mazza et al. (2012b, p. 110), who Stratigraphic occurrence: Uncommon in the medioconstricta Subzone emphasized the sub-circular, strongly ornate platform of the species. through spenceri Subzone of the samueli Zone. Nevertheless, included in this concept was an element with a more Carnepigondolella zoae (Orchard) rectangular platform (Mazza et al., 2010, pl. III, figs. 5a-c, sample Figure 35. 7-15 NA48), and one small specimen illustrated here shows some posterior rounding. It is unclear whether the species spatulata includes both 1991 Metapolygnathus n. sp. F - Orchard (a), p. 176, pl. 1, figs. varieties because no fully rounded morphotypes have been recovered 7-11; pl. 3, fig. B. from Black Bear Ridge. Based on their differing ages, the Canadian and 1991 Metapolygnathus zoae n. sp. - Orchard (b), p. 139, pl. 1, figs. Sicilian taxa may be homeomorphs. 7-9. 2012 [p] Carnepigondolella zoae (Orchard) central morphotype - Stratigraphic occurrence: Common in the spenceri Subzone of Mazza et al. (b), p. 103, pl. 4, figs. 1a-c (only). the samueli Zone in the Upper Carnian. Mazza et al. (2012b) record Epigondolella spatulata from the lower Lacian/ Lower Norian, above Holotype: GSC 95203, Figure 35. 13-15. the boundary interval, and well above the levels from which the Description: The P1 elements are relatively elongate with posterior Canadian elements occur. platforms that are initially narrow and later broaden and lengthen, Carnepigondolella spenceri sp. nov. becoming subquadrate with rounded posterior-lateral corners. They are Figure 38. 20-35 characterized by large, well defined but low nodes with a circular cross- section on each anterior platform margin. These margins are raised 2007 Carnepigondolella pseudoechinata (Kozur) - Orchard (b), relative to the flat posterior platform, and a submedial constriction fig. 1.4-6. separates the two parts. The free blade is about ⅓ element length and 2012 [?] Epigondolella cf. spatulata - Mazza et al.(b), p. 110, pl. 6, fig. has a relatively high convex profile; it descends onto the platform 9a-c. and continues as confluent nodes and finally an enlarged terminal Derivation of name: Named for Spencer, my grandson. node surrounded by a posterior platform brim. In adult forms like the holotype, the basal pit is situated a little posterior of the centre of the Holotype: GSC 132712, Figure 38. 24-26. platform, anteriorly shifted relative to the posterior keel. Type stratum: Bed 4, within the Pardonet Formation. Remarks: The original description of this species noted the position Age: The spenceri Subzone of the samueli Zone. of the pit near the posterior end of the narrow keel but noted that this position was variable. The holotype of Carnepigondolella zoae (re- Diagnosis: The P1 elements are characterized by a relatively short, illustrated here) from Haida Gwaii is larger than those recovered during equidimensional, subquadrate to subcircular platform, and a relatively this study, and has a pit that lies relatively farther to the anterior. The long blade equal to ~½ of the total element length. Particularly in present specimens from Black Bear Ridge have a pit that lies closer inner lateral profile, the lower surface of the posterior platform may be to the end of the keel and are regarded as earlier growth stages of the straight or even upturned in contrast to most contemporary species. The species. platform bears 2-4 small, mostly sharp anterior denticles, and a largely Noyan and Kozur (2007, p. 173-4) regarded Carnepigondolella unornamented posterior part that is laterally expanded with variably zoae as the appropriate name for Late Carnian (Tuvalian) elements rounded postero-lateral margins. A few isolated nodes may occur on the formerly assigned to “Metapolygnathus” nodosus, the type of which posterior margin. The carina is short and terminates at about the centre they thought was likely to be an older, Julian species. Based on their of the platform in a slightly larger discrete cusp, or rarely in a smaller holotypes, these two species clearly differ, and certainly none of the accessory node. The pit occupies a submedial position, and the keel North American Late Carnian specimens previously assigned to M. commonly bifurcates at its posterior end. nodosus by the present author are examples of C. zoae. Most are herein Comparisons: The platform of this species is relatively much shorter assigned to new species of Quadralella. and the blade is longer than older representatives of Carnepigondolella. Mazza et al. (2010, 2012b) included a broad morphological range Epigondolella heinzi Mazza, Cau and Rigo has a similar carina, pit, of specimens in this species and differentiated a Morphotype A and B in and keel, but it has a shorter free blade and a more ornate posterior addition to a central morphotype. Only the latter is clearly an example platform, which is an uncommon feature in C. spenceri (e.g., Fig. 38. of this species in this author’s view: the other morphotypes lack a 29-31). Both E. spatulata, and Carnepigondolella aff. spatulata of this platform constriction, have more rounded posterior outlines, and more variable anterior nodes, features that suggest affinity with Primatella 55 rotunda sp. nov. (q.v.). Carnepigondolella as the ancestor of Epigondolella but did not demarcate the two genera in the same way. Kozur (2003) described Stratigraphic occurrence: At its type locality, the species is not the oldest species as Epigondolella orchardi, which he regarded independently dated, but nearby on Kunghit Island, the species occurs as transitional between Carnepigondolella pseudodiebeli and above Welleri Subzone II and below Macrolobatus Zone ammonoid Epigondolella quadrata, but Mazza et al. (2010, 2012a, b) assigned collections, within the Late Carnian. At Black Bear Ridge, the species is E. orchardi to Carnepigondolella, and introduced an additional uncommon in the zoae Subzone through the medioconstricta Subzone transitional species, Epigondolella miettoi, between C. orchardi and E. of the samueli Zone. In Sicily, the various morphotypes of the species quadrata. The phyletic relationships proposed here differ (Figs. 18, 22). have the same stratigraphic range in the upper Tuvalian (Mazza et al., At Black Bear Ridge, Epigondolella aff. miettoi is recognized 2012b). as a separate taxon within the quadrata Zone but stratigraphically Carnepigondolella? sp. nov. A earlier examples are unknown. The species orchardi is here referred Figure 39. 21-23 to Primatella and is regarded as one of a spectrum of taxa ancestral to Epigondolella, but unrelated to C. pseudodiebeli. In this work, Description: A small P1 element with an elongate triangular, posteriorly Quadralella rather than Carnepigondolella is regarded as precursor to pointed platform, with sharp, irregularly developed anterior marginal Primatella. denticles, a short blade bearing 5 denticles, a carina that extends close to the posterior end of the platform, a centrally located pit, and a pointed Composition: The scope of this work does not extend above the lowest posterior keel. part of the quadrata Zone so only those Epigondolella taxa present in the lowest meter are considered here. Six species and two morphotypes Comparisons: This element is superficially similar to Acuminatella of the name-giver are recognized, most of which were formerly acuminata, but elements of comparable size differ: those of the latter combined as the E. quadrata population (Orchard, 1983). have a clearly longer and abruptly narrower posterior platform, and a relatively longer blade with more denticles (6-8). Stratigraphic occurrence: The first representatives of Epigondolella appear in the late Kerri Zone of the Lower Norian in several localities Remarks: This unique P1 platform element differs from those of in British Columbia (Orchard, 1983, 1991b). The genus thereafter contemporaneous Carnepigondolella species in the long carina dominates most collections through the remainder of the Lower and pointed platform and keel, morphological features that were Norian. Records of Epigondolella species in the Upper Carnian of excluded in the original diagnosis of the genus (Kozur, 2003). The Pizzo Mondello are anomalous in terms of the Canadian succession: specimen appears homeomorphic with Acuminatella, but because of they may be examples of Carnepigondolella (e.g., C. pseudodiebeli its stratigraphic position and denticle characteristics, an affiliation with beta morphotype). Carnepigondolella is suggested. Epigondolella aff. miettoi Mazza, Cau, and Rigo Stratigraphic occurrence: Rare in the zoae and ?medioconstricta Figure 40. 1-9 subzones of the samueli Zone. 2012 [aff.] Epigondolella mietto sp. nov. - Mazza et al. (a), p. 18, 19, Epigondolella Mosher 1968 fig. 9F-H Figures 22-25; 40, 41 2012 [aff.] Epigondolella mietto Mazza, Cau and Rigo, in press - Mazza 1968 Epigondolella n. gen. - Mosher (a), p. 935-36. et al. (b), 104, 106, pl. 5, fig. 1. 1991 Epigondolella Mosher – Orchard (b), p. 306. Description: An Epigondolella species with a relatively short, 2013 Epigondolella Mosher - Orchard, p. 449. subrectangular platform with a length-to-breadth ratio of between 1:1 Type species: Polygnathus abneptis Huckriede 1958 and 2: 1, high, apically sharp marginal denticles on the anterior margins only, a carina that terminates in a central cusp, and a submedial pit. Description: The P1 platform element has high and sharp anterior platform denticles that almost double the height of the relatively flat Remarks: These elements co-occur with Epigondolella quadrata at platform: when viewed in profile, the denticles are commonly between Black Bear Ridge but differ from that species in possessing a carina that about 35% and 45% of the total depth of the element measured from a terminates in a large cusp rather than in an additional node posterior lateral denticle tip to the basal keel margin. The platform basal margin of it. The same feature led Mazza et al. (2012a, p. 19) to establish E. is often straight or posteriorly stepped-up. Platform microreticulation miettoi, which they regarded as ancestral to E. quadrata. However, the is less compact and more subdued than in older genera, and it does present species differs from E. miettoi in its higher and sharper anterior not extend onto the platform denticles (Orchard 1983, fig. 9). The denticles, the generally longer free blade, and the upward arching of posterior platform may be unornamented or have weak-to-strong the lower surface. These are features that also serve to differentiate marginal platform nodes. The free blade is about one-third total element Carnepigondolella from Epigondolella, and to which E. miettoi sensu length and descends onto the platform as a carina that terminates in a stricto may belong, as suggested by its stratigraphic position. E. aff. central node, or continues onto the posterior platform as one or two miettoi differs from E. quadrata in having a shorter platform lacking additional nodes. The pit is submedial, anterior within a keel that is pronounced postero-lateral expansion. often bifurcated posteriorly, particularly in large specimens. Stratigraphic occurrence: Abundant at the base of the quadrata Zone Comparisons: Primatella P1 elements have lower, often blunter at Black Bear Ridge, but it is not yet determined how high it ranges. anterior denticles, a basal platform profile that is invariably downturned Mazza et al. (2012b, p. 106) recorded E. miettoi sensu stricto entirely in posteriorly, and a denser microreticulation that covers both the platform the uppermost Tuvalian. and anterior nodes (Orchard 1983, figs. 3A, B, F, E). Apically pointed Epigondolella quadrata Orchard anterior denticles of P1 elements of Carnepigondolella are typically Figure 40. 10-18; 19-27 more numerous and much smaller than in Epigondolella. The pit in both Primatella and Carnepigondolella often lies more to the posterior 1983 Epigondolella abneptis subsp. A population - Orchard, p. than in Epigondolella, although this is not the case in all species. A bifid 179-181, fig. 4 (part). keel may occur in each of these genera, and is not in itself regarded as 1991 Epigondolella quadrata n. sp. - Orchard (b), p. 311, pl. 2, a basis for separating them. figs. 1-3, 7-9. Remarks: All the ornate platform taxa of the Upper Carnian and Norian Holotype: GSC 95265, Figure 40. 16-18 were formerly referred to Epigondolella, particularly to the type species Diagnosis: P1 elements have a subrectangular to posteriorly expanded E. abneptis, which had an alleged range from the Late Carnian through platform with a length-to-breadth ratio of between 1.5:1 and 2:1, and much of the Norian. Orchard (1991a, c) distinguished some of these as a blade that is between ⅓ and ½ total element length; thin flanges may new Carnian species that he provisionally assigned to Metapolygnathus, extend close to the anterior end of the blade. A medial constriction may and which were later assigned to Carnepigondolella (Kozur, 2003). occur on one or both lateral margins, beyond which the platform may Epigondolella primitia was also assigned to Metapolygnathus based maintain uniform breadth or expand with acutely angled postero-lateral on its anterior pit, but now forms the basis for Primatella Orchard. corners. There are 2-3 high and sharply terminated anterior denticles The first Epigondolella species are regarded here as represented by E. on each lateral margin, the posteriormost of which is often the highest. quadrata and its associates, which first occur high in the Kerri Zone. The carina often ends in a prominent node, or a smaller accessory node Both Kozur (2003) and Mazza et al. (2012a) regarded 56

Figure 40. 1-9. Epigondolella aff. miettoi Mazza, Cau and Rigo. 1-3. GSC 132728, sample 32. 4-6. GSC 132729, sample 10/07. 7-9. GSC 132730, sample 32. 10-18. Epigondolella quadrata Orchard 1991, alpha morphotype. 10-12. GSC 132731, sample 32. 13-15. GSC 132732, sample 10/06. 16-18. GSC 95265, sample PH-213b. Holotype. Juvavites Cove. 19-27. Epigondolella quadrata Orchard 1991. beta morphotype. 19-21. GSC 132733, sample PH-213b. Juvavites Cove. 22-24. GSC 132734, sample BH-48. Brown Hill. 25-27. GSC 132735, sample 10/05. Scale bar = 200 microns. to its posterior, but neither one is the cusp; a broad posterior platform large posterior carinal nodes. As pointed out by Mazza et al. (2012b), brim is developed. The posterior platform is generally inornate but may the pit typically underlies neither of these nodes, which therefore are bear an incipient marginal node. The pit is located beneath the platform not the cusp. At Black Bear Ridge, the posterior platform shapes of E. midpoint, and the posterior keel usually bifurcates immediately quadrata form the basis for differentiating two morphotypes. posterior of the pit or a short distance beyond it. Stratigraphic occurrence: The elements assigned to this species Comparisons: Epigondolella aff. miettoi differs in its shorter platform appear, by definition, at the base of the quadrata Zone. The type and in having a terminal cusp rather than accessory posterior carinal stratum and locality of Epigondolella quadrata is at Juvavites Cove, nodes. Other species of Epigondolella that appear concurrently with where it is associated with ammonoids of the high Kerri Zone of the E. quadrata differ in having rounded, pointed, or ornate posterior Lower Norian. Mazza et al. (2012b, p, 108) questionably record both outlines, although they all have in common the characteristic high this species and E. miettoi first from the uppermost Tuvalian (Carnian) anterior platform denticles. The P1 elements of Primatella orchardi at Pizzo Mondello, although all their illustrations of E. quadrata and P. subquadrata, which have a carina that ends in a central cusp, and come from much higher Norian levels (samples NA 56, 58, 60) where those of P. stanleyi and P. mersinensis, which have additional posterior “advanced specimens”, like the holotype, occur. These Late Carnian carinal nodes, represent homeomorphic couplets similar to E. aff. “Epigondolella” elements, which appear before M. communisti, have an miettoi and E. quadrata. The Primatella species differ in their lower, uncertain relationship to Early Norian species from British Columbia. less-developed anterior denticles, more downturned posterior profile, They may be homeomorphs. Certainly, use of E. quadrata sensu Mazza and dense microreticulation. et al. (2010) as an index for the base of the Norian is untenable. Remarks: P1 elements of Epigondolella quadrata have one or two 57 alpha morphotype Epigondolella aff. uniformis Orchard Figure 40. 10-18 Figure 41. 1-10 1991 [p] Epigondolella quadrata n. sp. - Orchard, p. 311, pl. 2, figs. 1991 [aff.] Epigondolella triangularis uniformis n. subsp. - Orchard 1-3 (only). (b), p. 315, pl. 3, figs. 1-3. 2012 [?] Epigondolella quadrata Orchard - Mazza et al.(b), p. 106-08, Description: An Epigondolella in which the P1 platform elements are pl. 5, figs. 3a-c (only). relatively narrow with subparallel margins and a rounded, oblique, or Remarks: This morphotype corresponds to the holotype of the species truncated posterior margin. There are 2-3 large and upright denticles on in having an expanded posterior platform margin with acutely angled each anterior platform margin, and less pronounced, obliquely oriented postero-lateral corners. The element from Pizzo Mondello, tentatively marginal denticles on the posterior platform. A large, usually terminal included here, is an early growth stage of the species, but other carinal node is surrounded by a broad platform brim. The keel extends elements illustrated by Mazza et al. (2012b) are examples of the beta further to the posterior than the centrally located pit, and bifurcates at morphotype. its end. Stratigraphic occurrence: Abundant in the basal quadrata Zone. At Remarks: Orchard (1991b) introduced Epigondolella uniformis first, this morphotype is strongly subordinate to the beta morphotype, as a subspecies of E. triangularis Budurov to accommodate highly but it becomes relatively more common in the youngest beds sampled. ornate elements that lacked the strong postero-lateral expansion and the secondary carina that characterized the nominal subspecies. The beta morphotype younger holotype has a somewhat rounded posterior margin and, Figure 40. 19-27 although this character did not feature in its diagnosis, subsequent 1983 Epigondolella abneptis subsp. A - Orchard, p. 179-81, fig. authors have stressed the rounded margin as diagnostic for the species 15F. (Mazza et al., 2012b). The present material has less robust posterior 1991 Epigondolella quadrata n. sp. - Orchard (b), p. 311, pl. 2, ornament than the holotype of E. uniformis and so an aff. assignment figs. 7-9. is preferred. Similarly ornate E. vialovi Buryi and E. praetriangularis 2012 Epigondolella quadrata - Mazza et al. (b), p. 106-08, pl. 5, Kozur and Moix (in Moix et al., 2007) are shorter and posteriorly figs. 2, 4-10. expanded. 2013 Epigondolella quadrata Orchard - Carter and Orchard, Fig. Stratigraphic occurrence: Common from the base of the quadrata 7. 22-24. Zone, within the upper Kerri Zone. The holotype of E. uniformis came Remarks: This morphotype has a subrectangular posterior platform from the Dawsoni Zone of Juvavites Cove. with subparallel lateral margins. Some elements show some postero- Epigondolella aff. vialovi (Buryi) lateral expansion on one or both sides, but this is far less than that Figure 41. 11-19 which characterizes Epigondolella rigoi Noyan and Kozur. The elements illustrated by Mazza et al. (2012b) from Pizzo Mondello are 1989 [aff.] Metapolygnathus vialovi sp. nov. - Buryi, p. 44-45, pl. 5, all relatively squat elements like those originally illustrated by Orchard figs. 1-8, pl. 6, figs. 5-10. (1983) as E. abneptis subsp. A, one of which was referred to E. rigoi 1997 [aff.] Metapolygnathus vialovi Buryi - Buryi, pl. II, figs. 1-3, by Mazza et al. (2012a); however, it is doubtfiul whether that species 6-11. occurs in British Columbia. Description: The P1 platform element is subrectangular, broadening Stratigraphic occurrence: Abundant in the basal quadrata Zone, towards the posterior. Two to four large and broadened upright where it is the most common element present. denticles occur on each anterior margin, whereas smaller, lower, and obliquely oriented denticles occur in the posterior part. The free blade Epigondolella aff. stefanionensis Noyan is relatively short, comprising ⅓ of the element length. The posterior Figure 41. 20-34 carina generally ends well in front of the posterior margin, but one or 1983 Epigondolella abneptis subsp. A - Orchard, p. 179, 181, fig. more nodes may occur to the posterior. On the underside, a submedial 4L. pit lies close to the bifid posterior keel margin. 2007 [aff.] Epigondolella stefanionensis n. sp. - Noyan, in Noyan and Remarks: Several taxa resemble the present species, particularly Kozur, p. 169, fig. 4.5. Epigondolella vialovi, originally described from the Sikhote-Alin Description: The Epigondolella P1 platform elements are typically region of Russia (Buryi, 1989). The P1 elements from Black Bear slender with a rounded posterior margin that may become laterally Ridge correspond in most respects to the holotype of that species, expanded with growth. A large terminal cusp is surrounded by a broad but they have a more angular posterior outline and apparently higher platform brim. On each anterior lateral margin there are 2-3 high and and sharper nodes (no lateral view of the holotype was provided). sharp denticles, whereas the flat posterior platform is unornamented. A The platform shape is similar to E. aff. miettoi but that species lacks pit is located slightly posterior of mid platform, and the posterior keel significant posterior ornament. The platform is shorter and broader than is squared off or bifid. E. aff. uniformis. Epigondolella violiniformis Ji, Yang, Zong, and Wu is similar Remarks: Noyan (in Noyan and Kozur, 2007) introduced Epigondolella but that species has a more rounded posterior platform: this difference stefanionensis for elements from Greece that were compared with needs to be assessed as potentially intraspecific. Epigondolella some elements included in the E. abneptis subsp. A population by praetriangularis Kozur and Moix (in Moix et al., 2007) is even more Orchard (1983), but which differed from “typical” contemporaneous posteriorly expanded than E. violiniformis and, like that species, also E. quadrata in their longer platform and rounded posterior margin. The has subdued posterior ornament. It lacks the carinal alignment of Canadian specimens differ from E. stefanionensis in having much more the latter but, again, the significance of these variations is uncertain. prominent anterior denticles compared with the “short denticles or The long blade of E. praetriangularis serves to distinguish it from E. high nodes” of the latter, and in their more posteriorly upturned lower vialovi, although that of E. violiniformis is broken. See also Primatella profile. The present species also has 2-3 rather than the 3-4 anterior permica. denticles seen in the Greek species. Stratigraphic occurrence: Common at the base of the E. quadrata The older, and possibly related Primatella rotunda, is morphologically similar to Epigondolella aff. stefanionensis in its zone at Black Bear Ridge. Mazza et al. (2010, 2012b) record similar platform shape but the former has more numerous and lower denticles elements from the upper Tuvalian and into the Norian, which is the span and nodes in the carina, the blade, and on the anterior margins; also recorded for the type material of E. vialovi (Buryi, 1997). Epigondolella stefanionensis may be transitional between the two. Epigondolella sp. nov. A Stratigraphic occurrence: Common elements appear at the base of Figure 41. 35-37 the quadrata Zone at Black Bear Ridge. Similar Greek and Slovakian Description: Relatively squat P1 platform elements with a pointed specimens are also allegedly from the “quadrata Zone”, but are regarded posterior platform, two large denticles on each anterior lateral margin, as late Tuvalian in age (Noyan and Kozur, 2007). This emphasizes the and an additional small to incipient node on each posterior lateral misinterpretation of the quadrata Zone in some Tethyan sections. margin and at the posterior tip. The typical Epigondolella blade 58

Figure 41. 1-10. Epigondolella aff. uniformis Orchard. 1, 2. GSC 132736, sample 32. 3, 4. GSC 132737, sample 32. 5-7. GSC 132738, sample 32. 8-10. GSC 132739, sample 10/05. 11-19. Epigondolella aff. vialovi (Buryi). 11-13. GSC 132740, sample 10/07. 14-16. GSC 132741, sample 10/06. 17-19. GSC 132742, sample 10/06. 20-34. Epigondolella aff. stefanionensis Noyan. 20-22. GSC 132743, sample 10/06. 23-25. GSC 132744, sample 32. 26-28. GSC 132745, sample 32. 29-31. GSC 132746, sample 10/05. 32-34. GSC 132747, sample 32. 35-37. Epigondolella sp. nov. A. GSC 132748, sample 32. Scale bar = 200 microns. 59 descends into two small carinal nodes, including the cusp, followed like posterior margin distinguishes this species from all others. by a large terminal denticle at the center of the posterior platform. The Kraussodontus reversus lacks a free blade and has a narrower, more subcentral pit lies well anterior of the slightly bifid posterior keel. tapered platform without anterior geniculation points. K. peteri has a flattened posterior platform and a more prominent cusp. Most other Remarks: This uncommon species differs from most other Lower species of Kraussodontus have more anterior pits. Norian epigondolellids in its pointed platform, a feature resembling Middle Norian Mockina Kozur (1989b). Similarly small Primatella ex Stratigraphic occurrence: This species of Kraussodontus appears low gr. pseudoechinata differ in their more ovoid, less pointed platforms in the exposed Ludington Formation at Black Bear Ridge concurrently and smaller, less sharp anterior nodes. P. rhomboidale has a comparable with C. eozoae and is abundant through the eozoae-ludingtonensis platform node distribution but has smaller anterior nodes, and a more Subzone of the samueli Zone and into the sagittale-beattyi Subzone of posteriorly located pit. A direct derivation from the latter is plausible. the primitia Zone. These elements appear to retain an essentially juvenile morphology Kraussodontus margaretae sp. nov. common to Epigondolella species. Figure 42. 15-29 Stratigraphic occurrence: Present at the base of quadrata Zone. Derivation of name: Named for my mother, Margaret. Genus Kraussodontus Orchard Holotype: GSC 132758, Figure 42. 27-29. Figures 11, 12; 42-45 Type stratum: Bed 13a, within the Pardonet Formation. 2013 Kraussodontus gen. nov. - Orchard, p. 449, 450. Age: The angusta-dylani Subzone of the primitia Zone. Type species: Kraussodontus peteri Orchard, 2013. Diagnosis: The P1 elements are generally narrow and elongate with a Diagnosis: The P1 platform elements are characterized by straight length-to-breadth ratio of ~3:1, a slightly curved axis, and a posterior to curved, elongated or abbreviated ovoid platforms with largely margin that is truncated with rounded corners. The inner lateral margin subparallel lateral margins of generally uniform height, and a relatively is straight to slightly concave, whereas the outer margin is slightly rounded posterior margin of varying width. In some taxa, there is a convex. Platform margins are not conspicuously upturned, and all but medial to posterior platform narrowing on one or both sides; the the anteriormost platform appears relatively flat in lateral view: it bears posterior margin may be narrow and tapered, or broader and truncated, a broad zone of microreticulation but no marginal nodes. The platform but it is never broader than the anterior platform. The anterior margins margins may gently rise to the anterior, but there is no conspicuous are inornate to weakly ornate. Both relative blade length and pit position parapet formed. The free blade is of variable length, ranging between varies (Orchard, 2013). 1/5 and ¼ element length. The carina is variable, but in all elements Comparisons: Species of Kraussodontus are most similar to those terminates well in front of the posterior platform margin, where there is of Quadralella but they lack the typical quadrate outlines, commonly a wide brim. The cusp is inconspicuous, but the posterior carinal nodes expanded posterior platforms, strong lateral constrictions, reduced may be enlarged in smaller specimens. The pit lies beneath the center posterior platforms, and more ornate anterior margins. The rounded of the platform or slightly posterior of that point and is followed by a posterior platforms also contrast with those of Metapolygnathus broad, subrectangular posterior keel. species, which have a more anteriorly located pit than most species Comparisons: These P1 elements differ from those of Kraussodontus of the present genus. Species of Parapetella are distinguished by their ludingtonensis in bearing a flatter platform and medial pit. They are strongly differentiated anterior parapets, and those of Primatella by similar to those of K. rosiae, but the P1 element of that species has their more denticulate character. more upturned and ornate margins, particularly in the anterior part, and Remarks: Several morphological trends are recognized in this genus. a more rounded posterior margin. The primitive morphology consists of elements of uniform width, but Remarks: These are variable elements but they are united in their younger elements may show platform shortening or broadening, weak unornamented and relatively flat platforms. Younger examples have lateral constriction, or increased curvature. a shorter platform than older ones, and the oldest possible example, Composition: Eleven species and five variations or morphotypes are assigned to Kraussodontus aff. margaretae, has a different carina. differentiated in this new genus from Black Bear Ridge, only one of Stratigraphic occurrence: Uncommon in the angusta-dylani Subzone which resembles a named species, Kraussodontus reversus (Mosher). through the basal parvus Subzone of the primitia Zone. Stratigraphic occurrence: Kraussodontus first appears low in the Kraussodontus aff. margaretae samueli Zone of the Upper Carnian Ludington Formation at Black Bear Figure 42. 12-14 Ridge and ranges into the middle subdivision of the parvus Subzone. It is not known when the genus originated, but it is regarded as an offshoot Remarks: Two incompletely preserved elements resembling of Quadralella. It is known also from southeast Alaska (Katvala and Kraussodontus margaretae differ in the configuration of the posterior Stanley, 2008), and Japan (Koike, 1982). carina, which consists of two to three large nodes that are slightly offset form the main axis. Kraussodontus ludingtonensis sp. nov. Figure 42. 1-11 Stratigraphic occurrence: Rare in the zoae Subzone of the samueli Zone, and low in the sagittale-beattyi Subzone of the primitia Zone. Derivation of name: Named after the Ludington Formation, type locality Mount Ludington, northeastern British Columbia. Kraussodontus peteri Orchard Figure 43. 8-10; 11-16 Holotype: GSC 132750, Figure 42. 4-6 1982 [?] Neogondolella polygnathiformis (Budurov and Stefanov) - Type stratum: Bed 4, within the Pardonet Formation. Koike, p. 25-6, pl. 1, figs. 20, 21 (only). Age: The spenceri Subzone of the samueli Zone. 2008 Metapolygnathus polygnathiformis (Budurov and Stefanov) - Katvala and Stanley, fig. 39.7. Diagnosis: The P1 elements are characterized by an elongate 2008 Metapolygnathus sp. aff. M. nodosus (Hayashi) - Katvala platform with straight, subparallel and upturned lateral margins that and Stanley, fig. 39.30. have a slightly elevated, flat-topped profile, behind distinct anterior 2013 Kraussodontus peteri sp. nov. - Orchard, p. 451, fig. 3.22- geniculation points. The rounded posterior margin may be slightly 3.24. expanded laterally and is also upturned so as to produce a spoon- shaped morphology to the posterior platform. A free blade is about Holotype: GSC 132605, Figure 43. 8-10. ¼ to ⅓ total element length and bears about five denticles of uniform Type stratum: Bed 3, within the Pardonet Formation. height that descend onto the platform as a low carina that consists of 6-7 fairly discrete nodes, the central ones of which are smallest. The Age: The spenceri Subzone of the samueli Zone. inconspicuous cusp is surrounded by a wide posterior platform brim. Diagnosis: A species of Kraussodontus in which the P1 element has an The pit is terminal with respect to both the platform and the keel. oval platform of variable length with sub-parallel to biconvex margins Comparisons: The short, generally low blade and rounded, spoon- that taper to a narrowly rounded posterior outline. Platform margins are 60

Figure 42. 1-11. Kraussodontus ludingtonensis sp. nov. 1-3. GSC 132749, sample 0. 4-6. GSC 132750, sample 4. Holotype. 7, 8. GSC 132751, sample 2a. 9-11. GSC 132752, sample 0c. 12-14. Kraussodontus aff. margaretae. GSC 132753, sample 0. 15-29. Kraussodontus margaretae sp. nov. 15-17. GSC 132754, sample 16. 18-20. GSC 132755, sample 17c. 21-23. GSC 132756, sample 17a. 24-26. GSC 132757, sample 18e. 27-29. GSC 132758, sample 13a. Holotype. Scale bar = 200 microns. 61 upturned and slightly raised to the anterior where subdued nodes may species in lacking a free blade, are assigned to Kraussodontus reversus, occur, but the posterior platform is flat. The blade is typically short (¼ although the variability of the original species, from the Arctic, is total length), but appears relatively longer in younger representatives. uncertain. Mosher (1973) emphasized the reversal in width from small, The carina terminates in a conspicuous cusp that lies well in front of the medially widened specimens to medially narrower larger specimens, posterior margin and is surrounded by a broad brim. The pit lies near although this is not evident in the present material. Rather, the largest the posterior end of the platform and keel (Orchard, 2013). specimen from Black Bear Ridge lacks the medial narrowing that Comparisons: These elements most closely resemble Kraussodontus characterizes the holotype (re-illustrated by Orchard, 1991b), and also roberti, but they are distinguished by the more posterior position of has higher blade denticles. Nevertheless, that narrowing feature is the pit and, typically, a more narrowly rounded posterior platform. seen in other species of Kraussodontus, to which the species is newly The pits of K. rosiae and K. urbanae lay more anteriorward, and they referred. have a curved longitudinal axis. The older K. ludingtonensis has more Stratigraphic occurrence: Uncommon in the eozoae-ludingtonensis upturned margins throughout. Subzone through basal medioconstricta Subzone of the samueli Zone. Remarks: Two morphotypes are differentiated: alpha morphotype Kraussodontus roberti sp. nov. (Fig. 43. 8-10) is the holotype and stratigraphically oldest representative, Figure 43. 17-22; 23-40 and is characterized by a long platform and a relatively short blade; the younger beta morphotype (Fig. 43. 11-16) has a shorter platform and Derivation of name: Named for my brother, Robert. longer blade. Holotype: GSC 132770, Figure 43. 32-34 Stratigraphic occurrence: Both morphotypes are common, the alpha Type stratum: Bed 18f, within the Pardonet Formation. morphotype through the samueli Zone, and the beta morphotype from the spenceri Subzone of the samueli Zone into the sagittale-beattyi Age: Middle subdivision of the parvus Subzone of the primitia Zone. Subzone of the primitia Zone. Similar forms, illustrated only in upper Diagnosis: The P1 elements have a relatively elongate, flat platform view, occur in Keku Straight, southeast Alaska (Katvala and Stanley, with a straight axis, a length-to-breadth ratio of between 1.5:1 and 2008). The specimen from Japan combines a blade like the alpha 2:1, sub-parallel lateral margins, and a rounded posterior margin. The morphotype and a shorter platform like the beta morphotype. anteriormost platform margins of larger specimens may bear a few Kraussodontus praeangustus (Kozur, Mirauta and Mock) subdued nodes. A free blade equal to between ⅓-½ total element length Figure 44. 13-15 passes into a carina composed of low nodes of which the posteriormost 1 or 2 are often higher than those to the anterior. The pit is anteriorly 1980 Gondolella praeangusta n. sp. - Kozur, Mirauta and Mock (in shifted within the keel, occupying a position that is medial or a little to Kozur), p. 152. the posterior. 1980 Gondolella praeangusta Kozur, Mirauta and Mock - Kovacs and Kozur, pl. 11, figs. 5a-c. Comparisons: This species differs from Kraussodontus peteri in having a forward shifted pit, and a broader, less tapered posterior platform. Description: The P1 element has a narrow, uniformly tapered, pointed platform with slightly raised anterior margins, and pronounced anterior Remarks: Two morphotypes are differentiated: geniculation points. Short triangular denticles of the incomplete free alpha morphotype blade become progressively lower and pass into a carina that extends Figure 43. 23-40 to virtually the posterior end of the platform. The penultimate carinal node is a slightly enlarged cusp, which sits above a pit that is terminal Description: P1 elements with relatively long platforms and short within the keel. blades, subdued anterior nodes, or sometimes a single, relatively well developed node, are developed around the geniculation point. Comparisons: The illustrated element closely resembles the species originally described from Dobrogea, Romania, differing only in Remarks: Because of their relative size of the morphotypes, these its slight long-axis curvature and possibly larger blade denticles. elements might be later growth stages of the beta morphotype. However, The Romanian species is poorly known, but it is assumed that these the two do not have identical stratigraphic ranges. The alpha elements differences are intraspecific. Additional rare elements recovered during display some variation in pit position, with older representatives (Fig. this study (from Pardonet Hill east) share the same pointed platform 43. 23-25) having a pit that is less shifted to the anterior than the but differ in having an anterior pit like that of the successor species younger elements (Fig. 43. 26-28). Metapolygnathus angustus Kozur. However, that species differs in its Stratigraphic occurrence: Common in the sagittale-beattyi Subzone more quadrate platform and short carina. through the acuminata-prominens Subzone of the primitia Zone. Some less ornate elements of the pointed Acuminatella sagittale resemble the present species but their broader platform, anteriorly beta morphotype shifted pit, and ornate antero-lateral nodes serve to distinguish them. Figure 43. 17-22 These two taxa may have a common ancestor in Kraussodontus, as 2011 [?] Metapolygnathus praecommunisti sp. nov. - Mazza et al., p. might some Parapetella species (e.g., P. riteri), which differ in their 124-9, fig. 3B, D, E. strong parapets and short carinas. Description: Relatively small elements that have a subrounded to oval Stratigraphic occurrence: Rare in the acuminata-prominens Subzone platform with a length-to-breadth ratio of ~1.5:1, slightly raised and of the primitia Zone. “Gondolella” praeangusta was originally unornamented anterior margins, a free blade equal in length to the described from Julian strata in Romania. platform, a pit located under the center of the platform, and a short Kraussodontus reversus (Mosher) posterior keel. Figure 43. 1-7 Comparisons: It is not excluded that these elements are juveniles of Neogondolella the alpha morphotype, although the platforms are much shorter. These 1973 reversa n. sp. - Mosher, p. 169-70, pl. 20, figs. elements are also similar to Parapetella sp. nov. A but differ in the 9, 10, 13, 15-17. 1991 Metapolygnathus straight platform axis, more oval outline, and lack of well differentiated reversus (Mosher) - Orchard (b), p. 318, pl. parapets. 1, figs. 4-6. Description Stratigraphic occurrence: Very common in the angusta-dylani : Generally small, narrow and elongate P1 elements with a Subzone through the acuminata-prominens Subzone of the primitia biconvex platform that tapers evenly to a narrowly rounded posterior, Zone. The questionable elements from Pizzo Mondello and Pignola and anteriorly to the end of the relatively low fixed blade. The posterior in Italy included in Metapolygnathus praecommunisti by Mazza et al. end of small specimens displays slight indentations that appear lost (2011) occur in the Tuvalian. in larger specimens. The carina extends to close to the posterior end of the platform where the terminal cusp is conspicuous, inclined, and Kraussodontus rosiae sp. nov. surrounded by a narrow posterior brim. The pit is terminal with respect Figure 44. 1-12 to both the platform and the keel. Derivation of name: Named for Rosie Jassi in gratitude for her work Remarks: These specimens, which differ from most contemporary on the Triassic biostratigraphic database. 62 63 Figure 43 (facing page). 1-7. Kraussodontus reversus (Mosher). 1. Zone. GSC 132759, sample E. 2. GSC 132760, sample D. 3-5. GSC 132761, sample 0. 6, 7. GSC 132762, sample 0c. 8-10. Kraussodontus peteri Kraussodontus vancouverense sp. nov. Orchard, alpha morphotype. 8-10. GSC 132605, sample 3. Holotype. Figure 45. 1-21 11-16. Kraussodontus peteri Orchard, beta morphotype. 11-13. GSC Derivation of name: Named for the city of Vancouver, where this work 132763, sample 5. 14-16. GSC 132764, sample 3. 17-22. Kraussodontus was undertaken. roberti sp. nov., beta morphotype. 17-19. GSC 132765, sample 17c. 20-22. GSC 132766, sample 14b. 23-40. Kraussodontus roberti sp. Holotype: GSC 132785, Figure 45. 10-12. nov., alpha morphotype. 23-25. GSC 132767, sample 14b. 26-28. Type stratum: Bed 07-12c, within the Pardonet Formation. GSC 132768, sample 18. 29-31. GSC 132769, sample 13b. 32-34. GSC 132770, sample 18f. Holotype. 35-38. GSC 132771, sample 17b. 38- Age: The angusta-dylani Subzone of the primitia Zone. 40. GSC 132772, sample 18a. Scale bar = 200 microns. Diagnosis: The P1 elements have an hourglass shaped platform outline due to a submedial narrowing of the platform margins between laterally expanded anterior and posterior parts. The posterior platform has a Holotype: GSC 132775, Figure 44. 7-9. rounded, bulbous outline. The anterior platform may bear subdued Type stratum: Bed 17c, within the Pardonet Formation. nodes on the variably elevated margins. The free blade is ~⅓ element length but is flanked by platform flanges in large specimens. The carina Age: The acuminata-prominens Subzone of the primitia Zone. is composed of 5-6 nodes that are often highest at the posterior end. Diagnosis: The P1 elements have narrow elongate platforms with a A relatively wide posterior brim is developed. The pit is posterior of length-to-breadth ratio of ~3:1, a slightly curved axis, and a uniformly the midlength and in front of the squared-off or occasionally pointed rounded posterior margin. The inner lateral margin is straight to slightly posterior keel. concave whereas the outer margin is slightly convex, which produces a Comparisons: There are species of Parapetella and Primatella with platform of generally uniform width. The anterior platform margins are platform constrictions but they have more prominent anterior ornament. variably upturned and raised relative to the posterior platform, and bear incipient to moderately well formed, but coalescing marginal nodes. Stratigraphic occurrence: Common in the angusta-dylani Subzone The free blade is of variable length, commonly about ⅓ element length, into the middle subdivision of the parvus Subzone of the primitia Zone. and is composed of about 10 denticles of subequal height and width that Kraussodontus wendae sp. nov. descend onto the platform as a row of discrete nodes that extend close Figure 45. 22-32 to the posterior end of the platform. The posterior carinal node may be conspicuously larger than those to the anterior, or may be subequal in Derivation of name: Named for Wendy Squire in gratitude for her size. The pit lies posterior of the platform midlength and is followed by support during the course of this study. a short posterior extension of the keel. Holotype: GSC 132792, Figure 45. 30-32. Comparisons: These elements are similar to Quadralella Type stratum: Bed 07-18, within the Pardonet Formation. praecommunisti curvata, but they differ in having a posteriorly rounded outline rather than the rectangular platform of the latter. They differ Age: The acuminata-prominens Subzone of the primitia Zone. from Kraussodontus urbanae in being less curved and lacking the Diagnosis: P1 elements have a narrow, elongate platform with a length- posterior tongue-like extension. to-breadth ratio of ~3:1 and a relatively constant width throughout Stratigraphic occurrence: Very common in the angusta-dylani except for the slightly bulbous, incurved posterior ¼. This shape is Subzone through the lower subdivision of the parvus Subzone of the accentuated by slight constrictions on the inner and sometimes also on primitia Zone. the outer margins. The platform margin profile is flat or undulating, and the anterior margins are smooth or bear poorly developed nodes. Kraussodontus urbanae sp. nov. The blade is short, equal to ¼ the element length, and the carina is Figure 44. 16-27 long with 6-9 nodes reaching close to the rounded posterior margin; Derivation of name: Named for Kathleen Urban in gratitude for her a narrow platform brim is developed around the posterior 2-3 larger support during the course of this study. carinal nodes. The pit is located beneath the posterior platform, slightly displaced to the anterior within the broadened posterior keel. Holotype: GSC 132781, Figure 44. 25-27. Comparisons: These elements are much longer, narrower, and flatter Type stratum: Bed 18c, within the Pardonet Formation. than Kraussodontus vancouverense and have a posterior rather than a Age: Lower subdivision of the parvus Subzone of the primitia Zone. submedial platform constriction. Diagnosis: The P1 elements have a strongly curved platform with a Stratigraphic occurrence: Uncommon in the angusta-dylani Subzone length-to-breadth ratio of ~3:1, a distinctively convex outer margin, a through the acuminata-prominens Subzone of the primitia Zone. concave inner margin, and a rounded posterior margin; the posterior ½ Kraussodontus sp. nov. A of the platform is an incurved linguiform tongue. The raised anterior Figure 45. 33-35 margins bear weakly to moderately well-developed, round-topped nodes. The free blade is about ⅓ element length, and the carinal nodes Remarks: This unique specimen has a flat platform with a medial generally number 6 to 8 and extend close to the posterior platform indentation on its inner side, a slightly raised anterior margin devoid of termination, where they attain their maximum height; a platform nodes, a high arcuate blade, 4 carinal nodes that increase in size to the brim of variable width lies beyond. The pit is medial in position and posterior, an inconspicuous cusp, a posterior brim, a rounded posterior anteriorly displaced within the keel, which tapers to a narrow point near outline, and a submedial pit. It has an even flatter platform than K. aff. the inner postero-lateral margin. margaretae and K. wendae, although in common with the former it has one lateral platform indentation. Comparisons: These P1 elements are far more strongly curved than Kraussodontus rosiae and possess a distinctive linguiform tongue. Stratigraphic occurrence: Rare in the angusta-dylani Subzone of the Unlike the present species, the strongly incurved Parapetella clareae primitia Zone. has an expanded outer margin in front of the incurved tongue, as well Genus Metapolygnathus Hayashi 1968 as an unornamented parapet. Figures 13; 46-48 Remarks: The anterior platform margins of this species are moderately 1968 Metapolygnathus gen. nov. - Hayashi (a), p. 72. raised and resemble those of some Quadralella species; however, they 2012 Metapolygnathus Hayashi - Mazza et al.(b), p. 111-12. are retained in the present genus because of their affinity with other 2013 Metapolygnathus Hayashi - Orchard, p. 451-2. elongate and posteriorly rounded species. Type species: Metapolygnathus communisti Hayashi 1968. Stratigraphic occurrence: Common in the angusta-dylani Subzone through the lower subdivision of the parvus Subzone of the primitia Description: The P1 platform elements of this genus are subrectangular in outline, and have neither parapets nor strong nodes on the anterior 64

Figure 44. 1-12. Kraussodontus rosiae sp. nov. 1-3. GSC 132773, sample 17c. 4-6. GSC 132774, sample 18e. 7-9. GSC 132775, sample 17c. Holotype. 10-12. GSC 132776, sample 17b. 13-15. Kraussodontus praeangustus (Kozur, Mirauta and Mock). GSC 132777, sample 17b. 16-27. Kraussodontus urbanae sp. nov. 16-18. GSC 132778, sample 17b. 19-21. GSC 132779, sample 07-13a. 22-24. GSC 132780, sample 17c. 25-27. GSC 132781, sample 18c. Holotype. Scale bar = 200 microns.

Figure 45 (facing page). 1-21. Kraussodontus vancouverense sp. nov. 1-3. GSC 132782, sample 18a. 4-6. GSC 132783, sample 18f. 7-9. GSC 132784, sample 6. 10-12. GSC 132785, sample 14b. Holotype. 13-15. GSC 132786, sample 14a. 16-18. GSC 132787, sample 07-12c. 19-21. GSC 132788, sample 14b. 22-32. Kraussodontus wendae sp. nov. 22, 23. GSC 132789, sample 14a. 24-26. GSC 132790, sample 17c. 27-29. GSC 132791, sample 14a. 30-32. GSC 132792, sample 07-18. Holotype. 33-35. Kraussodontus sp. nov. A. GSC 132793, sample 15. Scale bar = 200 microns. 65 66 platform margins; low nodes may be present on one or both sides stratigraphically restricted, so they are identified here as different at a geniculation point. The free blade is between 1/4 and 1/2 element morphotypes. The oldest representative appears about half way length and extends onto the platform as a row of nodes of variable size through the exposed Ludington Formation and is distinguished from and spacing; the cusp may be terminal or be followed by additional, otherwise similar and contemporaneous Quadralella species by its sometimes larger nodes. The carina terminates in front of the posterior medial pit. Younger examples have slightly elevated anterior platform platform and is separated from it by a brim of variable width. Compact margins and a pit that is increasingly shifted anteriorly. The derivative microreticulae cover the platform margins. The pit is medial to anterior Metapolygnathus parvus has a strongly reduced and smooth platform, in position, and the keel extends far to its posterior where its termination longer free blade, and far anterior pit. Metapolygnathus dylani is squared-off or bifurcated. represents an intermediate form within this transition. The platform margin of the holotype of Metapolygnathus Comparisons: These elements differ from those of Primatella and communisti has a profile that gently rises to the anterior before Quadralella in lacking significant anterior marginal nodes, common descending to meet the blade. In contrast, Parapetella irwini has a platform constrictions, and/ or posterior pits. Parapetella differs in its conspicuously raised anterior margin, as does at least one specimen strongly differentiated anterior parapets. P1 elements of Kraussodontus illustrated by Mazza et al. (2012b, pl. 8, figs. ?5, 6). Specimens of M. are similarly unornamented but have narrower or rounded posterior communisti illustrated by Noyan and Kozur (2007) have generally outlines and/or posterior pits. Some stratigraphically young species planar lateral surfaces. of all the aforementioned genera have anteriorly located pits, which is no longer regarded as unique to Metapolygnathus. The older Remarks: Morphotypes of Metapolygnathus ex gr. communisti are genus Paragondolella has P1 elements with a flat platform without differentiated on the basis of platform outline and curvature: lateral geniculation points, a conspicuously high blade-carina, a posteriorly margins may be sub-parallel, biconvex, plano-convex, or concavo- located pit, and a different multielement apparatus (Orchard, 2005; convex. Stratigraphically younger morphotypes are more strongly contra Kiliç et al., 2013). downarched posteriorly and have the strongest long axis curvature. The pit is a little behind the platform center in the stratigraphically older Remarks: Orchard (1991a, c) referred all Upper Carnian and basal specimens, and a little in front of it in younger ones. Norian platform elements with quadrate outlines, geniculation points, Mazza et al. (2012b) most recently discussed Metapolygnathus and a free blade to Metapolygnathus, regardless of the pit position. This communisti and illustrated a growth series in which early representatives partly arose from the fact that an anterior pit and posterior prolongation had relatively short platforms and long blades (see also Krystyn, 1980). of the keel, essential features of the type species M. communisti, were These are also features of M. parvus, which these authors differentiated observed in several different lineages in which anterior migration also on the basis of its ovoid platform shape and lack of platform nodes. of the pit occurred over time. Assignment of all taxa with anteriorly The similarity had led authors to regard the two as synonyms, but at shifted pits to Metapolygnathus was judged to be more artificial than Black Bear Ridge the occurrence of M. parvus is much more restricted combining otherwise similar elements together until lineages could be stratigraphically. Notwithstanding the difficulty of distinguishing early better understood. Assignment of many Carnian taxa to Paragondolella, growth stages of these two species, I emphasize the relative blade and as has been common practice, is inappropriate because the latter genus platform lengths to differentiate them. is based on Middle Triassic P. excelsa, which differ as discussed above. Thus, the scope of both Metapolygnathus and Paragondolella Stratigraphic occurrence: Noyan and Kozur (2007) record all are restricted in this work. Only three species of the present genus are members of the Metapolygnathus communisti group as confined to the recognized, although these are grouped into ten morphotypes. latest Tuvalian, whereas Mazza et al. (2012b) record M. communisti from the latest Tuvalian to earliest Lacian. The collective range of Stratigraphic occurrence: At Black Bear Ridge, Metapolygnathus the morphotypes at Black Bear Ridge is much longer, from within ex gr. communisti and its derivatives range from midway through the the samueli Zone through most of the primitia Zone, which is in both samueli Zone to close to the top of the primitia Zone. the Upper Carnian and the Lower Norian. The stratigraphic range of Metapolygnathus ex gr. communisti Hayashi individual morphotypes is much less, and may point to their future Figure 46. 1-32 stratigraphic utility. 1968 Metapolygnathus communisti gen. et sp. nov. - Hayashi (a), p. Morphotype 1 72, pl. 3, fig.11a-c. Figure 46. 1-3 1980 Metapolygnathus communisti Hayashi morphotype A - Description: This unique specimen, the oldest representative of the Krystyn, pl. 12, figs. 8-14. group recovered, has the particular feature of downsloped anterior 2001 Metapolygnathus communisti Hayashi - Orchard et al. (a), pl. platform margins. It is more uniformly rectangular than younger forms, 4, fig. 7. with a straight inner margin and a slightly sinuous outer margin. The 2007 Metapolygnathus communisti communisti Hayashi - Noyan carina extends as a row of low nodes posterior of the small cusp. The pit and Kozur, p. 169, 171, figs. 7.3, 5-8. lays a little posterior of platform midlength, well in front of the forked 2010 Metapolygnathus communisti (Hayashi) - Orchard (a), fig. 13. 22, 23. posterior keel. 2012 [p] Metapolygnathus communisti growth series - Mazza et al.(b), Stratigraphic occurrence: Rare in the eozoae-ludingtonensis Subzone p. 112, 114, pl. 8, figs. 1-4, ?5 (only). of the samueli Zone. 2013 Metapolygnathus ex gr. communisti (Hayashi) - Carter and Morphotype 2 Orchard, fig. 13. 32-34. Figure 46. 4-6 2014 Metapolygnathus ex gr. communisti (Hayashi) - Balini et al., fig. 1.h1-3.1 2011 [?] Metapolygnathus praecommunisti sp. nov. - Mazza et al., p. Description: P1 elements included here are rather variable but are 124-9, fig. 3G (only). united in bearing elongate, subrectangular platforms with a length-to- Description: This morphotype has a platform that widens medially. breadth ratio of 2.5:1. Often, the outer platform margin is convex, the The anterior platform margins are level and then descend steeply to inner margin is straight to concave, and the posterior margin is straight the anterior. The free blade is relatively short and continues as a carina to obliquely truncate. The platform margins are generally of uniform composed of a row of rising nodes posterior to the small cusp. The pit height but may be slightly lower or slightly higher at the anterior end where one or two small marginal nodes may occur on one or both sides. The free blade varies between ¼ and ⅓ total element length Figure 46 (facing page). 1-32. Metapolygnathus ex gr. communisti and its denticles descend onto the platform as 3-6 variably developed Hayashi. 1-3. Morphotype 1. GSC 132794, sample D. 4-6. carinal nodes that extend beyond the inconspicuous cusp. A posterior Morphotype 2. GSC 132795, sample 1. 7-9. Morphotype 3. GSC platform brim of variable width is present. The pit is medial to anterior 132796, sample 92-3. 10-15. Morphotype 4. 10-12. GSC 132797, in position and far anterior of the posterior end of the keel, which may sample 6. 13-15. GSC 132798, sample 07-13a. 16-18. Morphotype 5. be bifurcated. GSC 132799, sample 17b. 19-24. Morphotype 6. 19-21. GSC 132800, sample 17b. 22-24. GSC 132801, sample 18a. 25-32. Morphotype 7. Comparisons: Seven variants of Metapolygnathus ex gr. communisti 25, 26. GSC 132802, sample 21d. 27-29. GSC 132608, sample 21a. 30- are recognized at Black Bear Ridge. None are common, but they are 32. GSC 132803, sample 30. Scale bar = 200 microns. 67 68 lies slightly posterior of platform midlength. The specimen figured from through asymmetrica-Norigondolella sp. subzones of the primitia Pizzo Mondello has anterior platform margins that decline gradually. Zone. Metapolygnathus communisti sensu Mazza et al. (2012b) appears Stratigraphic occurrence: Rare in the zoae Subzone of the samueli at a similar stratigraphic level to this morphotype in Pizzo Mondello. Zone. The Pizzo Mondello specimen comes from sample NA11, about Metapolygnathus dylani sp. nov. 11 m below the boundary interval. Figure 47. 1-21 Morphotype 3 2011 [p] Metapolygnathus praecommunisti sp. nov. - Mazza et al., p. Figure 46. 7-9 124-9, fig. 2H, I (only). Description: This element differs from other morphotypes in having 2012 [?] Metapolygnathus communisti Hayashi - Mazza et al. (b), p. a higher, crest-like free blade and a carina that terminates at a distinct 112, 114, pl. 8, fig. 5 (only). central cusp. A small node anterior of the cusp represents the end of a Derivation of name: Named for my grandson, Dylan. fixed blade that rises progressively to the anterior; a small accessory node occurs on the broad posterior platform brim. The platform Holotype: GSC 132808, Figure 47. 13-15. margins undulate but are essentially concave on the inner side, and Type stratum: Bed 14b, within the Pardonet Formation. straight to convex on the outer margin. The pit lies slightly posterior to platform midlength. Age: The angusta-dylani Subzone of the primitia Zone. Stratigraphic occurrence: Rare in the spenceri Subzone of the Diagnosis: The P1 element has a relatively short platform with a samueli Zone. length-to-breadth ratio of 2:1. Early growth stages have a narrowing of the platform at midlength, but larger elements have more or less straight Morphotype 4 lateral margins; the posterior margin is straight to oblique and has Figure 46. 10-15 variably rounded postero-lateral corners. The anterior lateral margins 2013 Metapolygnathus communisti Hayashi - Carter and Orchard, are generally raised compared to the posterior part, although not fig. 3. 32-34. abruptly, and rarely bear incipient nodes. The blade is equal to about ½ element length, and the carina is composed of 3-5 nodes that terminate Description: This morphotype has a narrower platform than the older in a large node separated from the posterior margin by a distinct brim, representatives and anterior platform margins that are raised and especially in larger specimens. The pit is submedial under the platform, developed into a single distinct node on one margin; incipient nodes may and anteriorly shifted in the keel, which has a squared-off termination occur on the opposite margin. The inner platform margin is concave, that reflects the shape of the platform margin. and the outer is convex, or both may undulate. The carina is composed of nodes of low to moderate height, and the cusp is terminal or followed Comparisons: These elements differ from Metapolygnathus ex gr. by several partly fused nodes. The pit lies at platform midlength. communisti in their relatively shorter platforms and longer blades. In this respect, they are intermediate between the former and M. parvus. Stratigraphic occurrence: Uncommon in the sagittale-beattyi through The platform is shorter than that of Quadralella praecommunisti, and angusta-dylani subzones of the primitia Zone. An example of this the anterior platform margins are neither as prominent nor as ornate. morphotype occurs at Sadler Point on Haida Gwaii, where it also occurs beneath parvus Subzone indicators (Carter and Orchard, 2013). Remarks: These elements exhibit variation in pit position and in their platform lateral profile: one specimen (Figure 47. 19-21) has Morphotype 5 a downsloped anterior lateral margin, similar to morphotype 1 of Figure 46. 16-18 Metapolygnathus ex gr. communisti. Further platform reduction in the Description: This unique element has a platform shape that differs group is believed to have led to the descendant M. parvus. A single from other morphotypes in being biconvex and having longer anterior specimen illustrated by Mazza et al. (2012b) has similar platform trough margins. The geniculation points are slightly raised but are dimensions to one example of this species (Fig. 47. 10-12) although the inornate. The carina extends close to the posterior end of the platform, former has a pit in a more anterior position. The specimens, regarded far posterior of the indistinct cusp. The pit lies anterior to platform as “advanced forms” of “M.” praecommunisti by Mazza et al. (2011), midlength. have a shorter platform than is typical of that species and are included here. Stratigraphic occurrence: Rare in the acuminata-prominens Subzone of the primitia Zone. Stratigraphic occurrence: Very common in the angusta-dylani Subzone of the primitia Zone. The specimen included here from Pizzo Morphotype 6 Mondello and identified aspraecommunisti (Mazza et al., 2011) comes Figure 46. 19-24 from the upper Tuvalian (sample NA19), whereas that referred to 2001 [?] Metapolygnathus communisti Hayashi - Orchard, in Orchard communisti comes from the boundary interval (sample NA37). et al. (a), Pl. 1, figs. 22, 23. Metapolygnathus ex gr. parvus Kozur Description: These are slender elements with a curved axis and Figure 48. 1-28; 29-48; 49-51 concavo-convex lateral margins. The slightly raised anterior platform 1972 Metapolygnathus parvus n. sp. - Kozur, p. 8, pl. 6, figs. 2-5. margins are inornate and terminate precipitously. The free blade may 2000 Metapolygnathus communisti (Hayashi) - Carter and Orchard, be longer than in older representatives and passes onto the platform as a pl. 1, figs. 4-7. strong carina that continues beyond the central cusp as a few fused and 2007 Metapolygnathus communisti parvus Kozur - Noyan and high nodes that may be separated from the cusp. The pit lies beneath the Kozur, p. 171, fig. 7.1. center of the platform or slightly to its anterior. 2012 Metapolygnathus parvus Kozur - Mazza et al. (b), p. 117, pl. Stratigraphic occurrence: Rare in the acuminata-prominens Subzone 8, figs. 9, 10. of the primitia Zone. A similar element was found in the Tezzeron Remarks: As discussed above, there has been some confusion Succession of the Cache Creek Complex in Nechako, B.C. (Orchard around the identity of Metapolygnathus parvus. The holotype has et al., 2001a). an unornamented ovoid platform, a long blade equal to half the total Morphotype 7 element length, and an anteriorly located pit, whereas small growth Figure 46. 25-32 stages of M. communisti allegedly are similar but with more rectangular platform shapes and an anterior node (as in M. echinatus Hayashi). In Description: Combined here are the youngest elements of the group, the present material from Canada, elements that share the characteristics which have a pit that is clearly located beneath the anterior half of of a long blade, an anterior pit, and generally unornamented platforms the platform. They include elements with straight to curved axes, and include both rectangular and more rounded platform outlines. Very few straight to concavo-convex lateral margins. The anterior platform if any of them conform exactly to the holotype of M. parvus, and yet margins may be slightly raised, and in one specimen the posterior their general morphology and stratigraphic occurrence suggests affinity platform is strongly downturned. Lateral margins may bear an incipient with the former species. Three morphotypes are differentiated here. node at the geniculation point. Stratigraphic occurrence: Metapolygnathus parvus appears at the Stratigraphic occurrence: Rare in the upper subdivision of parvus base of the parvus Subzone of the primitia Zone at Black Bear Ridge, 69

Figure 47. 1-21. Metapolygnathus dylani sp. nov. 1-3. GSC 132804, sample 14b. 4-6. GSC 132805, sample 14b. 7-9. GSC 132806, sample 14a. 10-12. GSC 132807, sample 14b. 13-15. GSC 132808, sample 14b. Holotype. 16-18. GSC 132809, sample 14b. 19-21. GSC 132810, sample 14b. Scale bar = 200 microns. and its range defines that subzone. The species occurs at the base of the Stratigraphic occurrence: Abundant in the parvus Subzone of the “boundary interval” at Pizzo Mondello (Mazza et al., 2012), but ranges primitia Zone. a little higher than that interval. beta morphotype alpha morphotype Figure 48. 29-48 Figure 48. 1-28 2007 [p] Metapolygnathus echinatus (Hayashi) - Orchard (c), pl. 1, 2007 Metapolygnathus echinatus (Hayashi) - Orchard (c), pl. 1, figs. 10-12, 22-24 (only). figs. 19-21 (only). Description: The P1 elements are generally small with a relatively 2010 Metapolygnathus parvus Hayashi - Orchard (a), fig. 13. 24. narrow, elongate, sub-rectangular platform with a length-to-breadth 2013 Metapolygnathus parvus Hayashi - Carter and Orchard, fig. ratio of between 2:1 and 3:1. Lateral platform margins are sub-parallel 3. 35-37. or slightly curved and the posterior margin is often squared-off with Description: The P1 elements are generally small and have a platform rounded corners. The platform margins are slightly raised to the that is sub-rectangular with a length-to-breadth ratio of between 1:1 anterior but bear no nodes. The free blade is at least ½ total element and 1.5:1. Lateral platform margins are sub-parallel or slightly curved, length and relatively longer in small elements. There are 3-4 carinal and the posterior margin is often squared-off but may be obliquely nodes of variable size and fusion, and these are more conspicuous in truncated or rounded. The platform margins are generally slightly small elements in which the platform is very narrow or even vestigial. raised to the anterior and may have a pronounced geniculation point, A posterior platform brim of variable width is developed. The pit but they rarely bear nodes, and nor is there a conspicuous parapet. characteristics are like that of the alpha morphotype. The free blade is at least ½ total element length, longer in small Remarks: This morphotype is relatively longer and narrower than the elements. There are 3-4 carinal nodes of variable size and fusion, alpha morphotype. Early growth stages have a very narrow platform which terminate in front of a posterior platform brim that is up to half that enlarges laterally during growth. the platform length. The pit lies beneath the platform-blade junction in small elements and behind this point in larger ones, but always Stratigraphic occurrence: Abundant in the parvus Subzone of the beneath the anterior half of the platform. primitia Zone. Remarks: Most elements have short subrectangular platforms like gamma morphotype the holotype of Metapolygnathus parvus, but many have irregular Figure 48. 49-51 margins. They differ from M. dylani in their shorter platform, longer 2007 Metapolygnathus n. sp. R - Orchard (c), pl. 1, figs. 4-6. blade, and more anterior pit. The beta morphotype has a relatively longer and narrower platform. Description: Small P1 elements that either lack a platform or have 70 71 Figure 48 (facing page). 1-28. Metapolygnathus ex gr. parvus Kozur, 10, 11. alpha morphotype. 1, 2. GSC 132811, sample 20f. 3-5. GSC 131170, 1982 [p] Neogondolella navicula (Huckriede) - Koike, p. 23-4, pl. 2, sample 20. 6-8. GSC 132812, sample 18h. Broken in lower view. 9-11. figs. 26, 27, 32, 33 (only). GSC 132813, sample 21b. 12, 13. GSC 132814, sample PH-3. Pardonet 1991 ‘Neogondolella’ navicula (Huckriede) - Orchard (a), pl. 4, Hill east. 14-16. GSC 132609, sample 18h. 17-19. GSC 132815, sample fig. 13. 18c. 20-22. GSC 132816, sample 18c. 23-25. GSC 132817, sample 2000 Norigondolella navicula (Huckriede) - Carter and Orchard, 18h. 26-28. GSC 132818, sample 20c. 29-48. Metapolygnathus ex gr. pl. 1, fig.9. parvus Kozur, beta morphotype. 29-31. GSC 132819, sample 18h. 2003 [p] Norigondolella navicula (Huckriede) - Channel et al., fig. A2. 32-34. GSC 132820, sample 20f. 35-37. GSC 131171, sample 20. 38- 7, 8, 10, 12, 13, 15, 21, 34; fig A3. 11 (only). 40. GSC 132821, sample 21. 41-43. GSC 131169, sample 20. 44-46. Description: The juvenile P1 elements have an incipient platform on GSC 132822, sample 20b. 47, 48. GSC 132823, sample PH-3. Pardonet the flanks of a blade bearing seven sharp, slightly inclined and discrete Hill east. 49-51. Metapolygnathus ex gr. parvus Kozur, gamma denticles, the posteriormost of which is a broader, subtriangular cusp morphotype. GSC 131167, sample PHE-23. Pardonet Hill east. Scale sitting above a terminal and expanded basal cavity. With growth, when bar = 200 microns. additional denticle are added to number 12 in total, the posterior carinal denticles are lower, blunter, and partly fused, whereas the anterior ones remain more discrete like those of the juveniles. Significantly, the a vestigial one represented by a thickening of the lateral flanks of a broader cusp becomes the penultimate carinal node and is followed by a blade-like element with ~10 blade denticles followed by commonly narrow but equally high and more discrete terminal denticle. The basal 3 lower carinal nodes above the inturned, initially upturned, and then cavity becomes relatively smaller with growth within a furrowed keel downturned posterior margin. In the absence of a platform brim, carinal that extends beneath the anterior fixed blade. Larger subadult elements denticles extend to the posterior end of the element. The flared margins show stronger upturning and thickening of the platform margins, a of the keel extend laterally beyond the margins of the element and largely fused posterior carina with an indistinct cusp, and a terminal contain a pit close to the carina-blade junction. denticle that is higher than those to its anterior. The anterior blade rises Comparisons: This form differs from early growth stages of above the anteriorly tapered platform as larger, partly fused triangular Metapolygnathus parvus beta morphotype in entirely lacking a platform denticles that form an low arched crest. Slight inturning of the posterior and having a more developed carina at its posterior end. platform may occur as that part of the platform becomes broader, and the margins of the basal cavity thicken producing a relatively smaller Remarks: This species is thought to be the end member of the lineage pit. Large adult elements have ~16 carinal denticles, half of which that includes Metapolygnathus parvus beta morphotype as its forbearer. are largely fused into a ridge, and these pass anteriorly into less fused This “naked gondolellid” formed the basis of earlier speculation but coalescing denticles that increase in size in that direction. Large (Orchard, 1991a) that Norigondolella may have been derived from platforms commonly have their maximum breadth at the posterior end, Metapolygnathus through loss of a posterior platform and subsequent but may be more biconvex, in common with earlier growth stages. A gain of a more extensive platform. However, the differing lower prominent posterior cusp is typically surrounded by a narrow platform surface morphology of the two argues against this, as does the fact that brim, although there is a trend for the cusp to become terminal. The Norigondolella is now known to appear earlier than M. parvus. pit and its surrounding keel occupy about one-half the breadth of the Stratigraphic occurrence: Abundant in the middle and upper element underside. subdivisions of the parvus Subzone of the primitia Zone. Comparisons: Norigondolella norica differs in being relatively less Genus Norigondolella Kozur arched with a much narrower platform, a higher blade-carina with more upright denticles, and a more expanded basal cavity than comparably 1990 Norigondolella n. gen. - Kozur, p. 127-131. sized elements of N. navicula. N.? hallstattensis has a much broader Type species: Paragondolella steinbergensis Mosher, 1968 medial platform and a higher blade. Remarks: This genus was introduced to accommodate Norian Remarks: This is the most common Norigondolella species pastiniplanate elements that had a depressed keel unlike the older encountered in this study, and is often the only one present. This has Neogondolella Bender and Stoppel, but similar to the more primitive facilitated confirmation of the growth series previously demonstrated Mesogondolella Kozur. The multielement apparatus is also known by others (e.g., Mosher, 1968b; Channel et al., 2003). Early growth to differ (Orchard, 2005). Norigondolella is common in most of the stages from the upper Lower Norian may be indistinguishable from samples from the Lower Norian Kerri Zone starting from bed 25 of those of N. hallstattensis; those illustrated by Koike (1982, pl. 2, figs. the Pardonet Formation at Black Bear Ridge (Fig. 3). Below that level, 28-31) could belong to either species. only single specimens are known in large conodont collections from Two morphotypes are differentiated based on the character of beds 13c, 17b, 18, 18e, 18f, and 18h, and they are too few to be sure of the posterior platform margin and cusp. The alpha morphotype their affinity. In the past, most of the younger occurrences have been corresponds to the well known P1 elements which have a subterminal, assigned to Norigondolella navicula (Huckriede) (Tables 6, 8), but two upright cusp and a distinct posterior platform brim, whereas the beta morphotypes of that species and one new species are distinguished morphotype has a more prominent, often more posteriorly projecting here. There are apparently no occurrences of these Norigondolella terminal cusp similar to that of the younger N. steinbergensis (Mosher). species at Pizzo Mondello, where both N. cf. navicula sensu Mazza The two species show similar variability in platform shape. et al. (2012b) from the Upper Carnian, and N. trinacriae Mazza, Cau Very rare examples of P1 elements of Norigondolella known from and Rigo from the Lower Norian differ in many respects. Neither of the boundary interval and older beds (13c, 17b, 18, 18e) appear to have these taxa is currently known from North America. In that connection, a more evenly tapered anterior platform than that seen in many later inclusion of the specimen of N. steinbergensis (Mosher) illustrated by examples of N. navicula. Also, the secondary denticle posterior of the Orchard (1991a) from Haida Gwaii in synonymy with N. cf. navicula cusp evident in juveniles of this species is seen for the first time in a (Mazza et al., 2012b, p. 122) is not supported. specimen from bed 18f. For now, the older elements are retained within the present species, pending more material from these levels. Stratigraphic occurrence: Traditionally regarded as Norian, although uncommon elements also occur in the Carnian (Nicora et al., 2007; Stratigraphic occurrence: Questionably and rarely in the angusta- Mazza et al., 2010, 2012b; herein). dylani Subzone through lower division of the parvus; rare in the middle division of the parvus Subzone, and abundant in the asymmetrica- Norigondolella navicula (Huckriede) Norigondolella Subzone of the primitia Zone. Within the latter subzone, Figure 49. 1-14. the alpha morphotype occurs first, and the beta morphotype become 1958 [p] Gondolella navicula n. sp. - Huckriede, p. 147-48, pl. 12, more common later. figs. 5, 6, 10 (only). Norigondolella norica sp. nov. 1968 Paragondolella navicula navicula (Huckriede) - Mosher (b), Figure 49. 15-19. pl. 119, figs. 11-23. 1973 Neogondolella navicula navicula (Huckriede) - Mosher, p. 1991 “Neogondolella” navicula (Huckriede) - Orchard (a), pl. 2, 168, pl. 20, figs. 11, 18. figs. 25-27. 1980 Neogondolella navicula (Huckriede) - Krystyn, pl. 11, figs. 2005 [?] Norigondolella steinbergensis (Mosher) - Orchard, p.89, 72

Figure 49. 1-8, 12-14. Norigondolella navicula (Huckriede) alpha morphotype. 1, 2. GSC 132824, sample 28. 3-5. GSC 132825, sample PHE- 26. 6-8. GSC 132826, sample 10/03. 12-14. GSC 132827, sample 25. 9-11. Norigondolella navicula (Huckriede) beta morphotype. GSC 132828, sample 10/03. 15-19. Norigondolella norica sp. nov. 15, 16. GSC 132829, sample PH316b. 17-19. GSC 131172, sample 26. Holotype. 20, 21. Misikella? sp. nov. A. GSC 132830, sample 07-13a. 22, 23. Misikella longidentata Kozur and Mock. GSC 132831, sample 04-115. Scale bar = 200 microns. figs. 15A-I of the platform; this has thickened margins rather than a platform 2007 Norigondolella navicula (Huckriede) - Orchard (c), pl. 2, surround. The lower profile of the element is relatively straight except figs. 16-18. for the posterior part, and it has a strongly flared basal cavity and 2010 Norigondolella navicula (Huckriede) - Orchard (a), figs. 13. anteriorly grooved keel. 4-6. Comparisons: This species resembles elements of N. steinbergensis Derivation of name: From its exclusive occurrence in strata of Norian from the Middle Norian (Orchard, 2005, fig. 15), but those specimens age. are more strongly arched, have a broader platform, and a relatively Holotype: GSC 131172, Figure 49. 17-19. larger cusp. See also Norigondolella navicula. Stratigraphic occurrence: Abundant in, and apparently restricted Type stratum: Bed 26, from within the Pardonet Formation. to, the lower part of the asymmetrica-Norigondolella Subzone of the Age: The asymmetrica-Norigondolella Subzone of the primitia Zone. primitia Zone. The same species occurs at a comparable level within Diagnosis: The P1 elements have a relatively narrow, flange-like the lower Kerri Zone at Juvavites Cove. platform bordering the margins of a high arcuate blade-carina composed Genus Parapetella Orchard of ~15 narrow, partly fused denticles that are highest at the anterior end Figures 14-16; 50-60 of the elements but which remain well above the level of the platform throughout its length. A triangular cusp is larger and more discrete than 2013 Parapetella gen. nov. - Orchard, p. 452. the anterior carinal denticles, and is followed by a prominent terminal Type species: Parapetella prominens Orchard, 2013. posterior denticle that is inclined above the downturned posterior end 73 Diagnosis: P1 platform elements of variable outline have distinctly have raised anterior margins, but they lack well differentiated parapets. raised anterior platform margins that are often differentiated into distinctive parapets that may be round-topped or angular in profile with Remarks: The anterior parapets in P1 elements of this genus substitute an anterior edge that descends to meet the blade gradually or abruptly for the denticles in Acuminatella and Primatella. Similar or even (Orchard, 2013). identical platform shapes found in these genera occur in Parapetella, which is regarded as having developed from inornate Quadralella Description: The platform shapes of the P1 elements vary considerably, species by progressive elevation of the anterior platform margins from relatively broad to narrow, and from relatively long to short; all and increasing differentiation of marginal parapets. Some species are are posteriorly downturned. Upper view outlines include those that are characterized by a medial constriction, others by posterior attenuation, subrectangular, oval, pointed, linguiform, medially constricted, and but both groups show anterior platform reduction through time (as posteriorly attenuated, expanded, or strongly reduced. In all cases, the in contemporaneous Metapolygnathus). A total of 17 species, plus a anterior platform margins are clearly higher (when viewed laterally) further seven unnamed, and seven morphotypes of Parapetella are than the posterior parts and increasingly differentiated so as to produce described here. high parapets or buttresses that are round-topped or angular in profile. The anterior edge of the platform descends to meet the blade gradually Stratigraphic occurrence: Several species of Parapetella make their or abruptly. Platform nodes occur rarely, although the parapets may be appearance at Black Bear Ridge at or near the base of the sagittale- weakly denticulate. Compact microreticulae cover both the platform beattyi Subzone of the primitia Zone. The youngest species occur in the asymmetrica-Norigondolella sp. Subzone of the primitia Zone and and parapets. The free blade is ⅓ to 2/3 total element length, depending on the extent of platform reduction. The carinal nodes may terminate in one is known from immediately beneath the Lower Norian quadrata front of a wide platform brim, or extend close to the posterior margin Zone. The genus occurs also on Haida Gwaii (Carter and Orchard, and be separated by a narrow brim: it may end in either a conspicuous 2013). cusp, or two or more fused nodes. A pit is posterior in early species, Parapetella beattyi sp. nov. medial in later ones, and anterior in the youngest. A posterior keel Figure 50. 1-18 may have a quadrate, pointed, or bifurcated termination, reflecting the platform shape. 2006 [?] Metapolygnathus n. sp. C - Orchard, pl.7, figs. 8-10. 2007 Metapolygnathus n. sp. Q - Orchard (c), pl. 1, figs. 1-8. Comparisons: Many species of Parapetella have a platform outline 2010 Metapolygnathus n. sp. Q - Orchard (a), fig. 13. 16. comparable to those of Primatella, but they have a parapet rather than 2013 Parapetella sp. - Carter and Orchard, figs 3. 26-28. strong anterior nodes or denticles. The weakly ornate or inornate genera Kraussodontus, Metapolygnathus, and Quadralella may sometimes Derivation of name: Named for Tyler Beatty, who ably assisted in the

Figure 50. 1-18. Parapetella beattyi sp. nov. 1-3. GSC 131240, sample 5. 4-6. GSC 132832, sample 5. 7-9. GSC 132833, sample 5. 10-12. GSC 131241, sample 5. 13-15. GSC 132834, sample 07-8. Holotype. 16-18. GSC 132835, sample 5. Scale bar = 200 microns. 74 field studies and conodont sampling. acuminata-prominens subzones, and ?basal parvus Subzone of the Holotype: GSC 132834, Figure 50. 13-15. primitia Zone. The species also occurs on Haida Gwaii where it and similar taxa with anteriorly shifted pits were the basis for records of Type stratum: Bed 07-8, from within the Pardonet Formation. Metapolygnathus communisti (Carter et al., 1989). Age: The sagittale-beattyi Subzone of the primitia Zone. Parapetella clareae sp. nov. Diagnosis: P1 elements have a sub-rectangular platform of variable Figure 51. 10-18 outline and an average length-to-breadth ratio of ~2:1. Many specimens 2011 [p] Metapolygnathus praecommunisti sp. nov. - Mazza et al.(a), have sub-parallel lateral margins that are indented on one or both sides p. 124-29, fig. 2, D?, E (only). in the posterior third; the largest specimens are of more uniform breadth. The posterior margins are generally straight but may be rounded postero- Derivation of name: Named for the late Clare Boggan, an inspiration laterally and occasionally asymmetric with one corner more expanded. during this study. In lateral profile, the anterior margins are raised, conspicuously in Holotype: GSC 132838, Figure 51. 10-12 smaller specimens in which the resulting buttresses are short and often very high. In larger specimens, they are longer and less differentiated. Type stratum: Bed 18a, within Pardonet Formation. Marginal ornament is often absent and, if present, restricted to a few Age: The acuminata-prominens Subzone of the primitia Zone. incipient nodes developed on the anteriormost platform. The blade is about ⅓ total element length and descends onto the platform as a row Diagnosis: The P1 elements have a strongly asymmetric platform of 4-6 variably spaced nodes that differ in size and fusion. Commonly, with an expanded, convex outer posterior outline and an extended the posteriormost two nodes are higher and partly fused, but often there inner postero-lateral lobe. The outer margin is initially straight but is a single large and discrete terminal cusp, or a smaller accessory node then expands outwards before curving inwards to form the posterior formed on the otherwise inornate posterior platform brim. The pit lies margin of the inner process. The inner margin is initially straight between the middle and posterior end of the platform, a short distance and then variably concave. The anterior margins are distinctly raised in front of the posterior end of the broad, squared-off keel. and generally inornate. The free blade is about ⅓ element length and continues as a row of closely spaced nodes that become more discrete at Comparisons: These elements differ from otherwise similar the point where the carina is deflected to the inner side. The posteriormost Quadralella species like Q. tuvalica by possessing clearly elevated and nodes are larger than those to the anterior and terminate either well in generally inornate anterior platform margins. The younger Parapetella front of the posterior end of the process, or close to its end; a brim of prominens may have a similar platform outline but has much more variable width is developed. The pit is located beneath the middle of strongly differentiated parapets in adult specimens, although early the platform where posterior expansion begins, and lies within a keel growth stages of the present species may have equally conspicuous that extends beneath the inner process and tends to bifurcate with a parapets. secondary keel extending towards the outer side expansion. Remarks: This species is the first to show marked differentiation Comparisons: These distinctive asymmetrical P1 elements resemble of anterior parapets and appears to have developed from species those of Kraussodontus urbanae, but that species is more anteriorly of Quadralella that are otherwise very similar in upper view. The ornate, and its more elongate platform has an evenly curved outer transition may have been gradual because, in some cases, the parapet margin rather than a posteriorly expanded one. is more developed on one margin: this is the case in the Yukon element assigned to Metapolygnathus n. sp. C (Orchard, 2006). Remarks: Elements such as these were included as “asymmetric morphotypes” of Metapolygnathus praecommunisti by Mazza et al. Stratigraphic occurrence: Abundant in the sagittale-beattyi Subzone (2011), but they are separated here because they have a more restricted through the basal part of the parvus Subzone of the primitia Zone. The occurrence than Quadralella praecommunisti subspp. (q.v.) sensu this species was also found in the Peril Formation of Haida Gwaii (Carter report. and Orchard, 2013), and the Yukon specimen was found in the Jones Lake Formation (Orchard, 2006). Stratigraphic occurrence: Common in the sagittale-beattyi Subzone through lower subdivision of the parvus Subzone of the primitia Zone. Parapetella broatchae sp. nov. Mazza et al. (2011, fig. 2E) figured a typical specimen of this species Figure 51. 1-9 from sample NA28 at Pizzo Mondello, which is ~13 m below the 1989 Metapolygnathus communisti (Hayashi) - Carter et al., pl. 1, boundary interval. An older element from sample NA14 (op. cit., fig. fig. 7. 2D) lacks the sinuous outer margin and may be a transitional form. 2007 [p] Metapolygnathus n. sp. Y - Orchard (c), pl. 2, figs. 37-39 Parapetella columbiense sp. nov. (only). Figure 52. 1-24 2013 Kraussodontus n. sp. aff. peteri Orchard - Carter and Orchard, fig. 3. 23-25. Derivation of name: Named for the province of British Columbia, in which Black Bear Ridge is located. Derivation of name: Named for Jane Broatch for her help in project microscopy. Holotype: GSC132855, Figure 52. 22-24. Holotype: GSC 132837, Figure 51. 4-6 Type stratum: Bed 17c, within Pardonet Formation. Type stratum: Bed 14b, within the Pardonet Formation. Age: The acuminata-prominens Subzone of the primitia Zone. Age: The angusta-dylani Subzone of the primitia Zone. Diagnosis: The P1 elements are characterized by a long and narrow tapered platform with a length-to-breadth ratio of 3+:1. These are Diagnosis: The P1 elements are relatively long and unevenly tapered to broadest near the anterior end, where the margins are subparallel or a narrow, roundly pointed posterior margin. The outer margin is broadly slightly expanded towards the posterior, and then taper strongly to the convex, particularly in the posterior half, whereas the inner lateral posterior from about mid-length to a narrowly rounded posterior end. margin is more or less straight or indented in the posteriormost 1/6 of the platform. The anterior margins are distinctly raised compared with the posterior margin and form long inornate parapets. The free blade is Figure 51 (facing page). 1-9. Parapetella broatchae sp. nov. 1-3. about ⅓ element length, and the carina continues as 4-7 nodes of which GSC 132836, sample 18. 4-6. GSC 132837, sample 14b. Holotype. the posteriormost one or two are usually the largest. The pit lies a little 7-9. GSC 131176, sample 16. 10-18. Parapetella clareae sp. nov. 10- posterior of the platform center and is anteriorly shifted within the keel, 12. GSC 132838, sample 18a. Holotype. 13-15. GSC 132839, sample which is squared off at its posterior termination. 18e. 16-18. GSC 132840, sample 14a. 19-27. Parapetella sp. nov. A. Comparisons: Similarly shaped, tapered platforms of Parapetella 19-21. GSC 132841, sample 5. 22-24. GSC 132842, sample 14b. 25- prominens angulare are shorter and have much more differentiated 27. GSC 132843, sample 04-16. 28-32. Parapetella sp. nov. B. 28-30. parapets. The platform outline mimics some Kraussodontus species but GSC 132844, sample 12a. 31, 32. GSC 132845, sample 12a. 33-38. those species do not have a parapet. Parapetella sp. nov. C. 33-35. GSC 132846, sample 17b. 36-38. GSC 132847, sample 17c. Scale bar = 200 microns. Stratigraphic occurrence: Common in the angusta-dylani through 75 76

Figure 52. 1-24. Parapetella columbiense sp. nov. 1-3. GSC 132848, sample 17b. 4-6. GSC 132849, sample 17b. 7-9. GSC 132850, sample 18d. 10-12. GSC 132851, sample 18d. 13-15. GSC 132852, sample 17c. 16-18. GSC 132853, sample 17c. 19-21. GSC 132854, sample 18d. 22-24. GSC 132855, sample 17c. Holotype. Scale bar = 200 microns. Long anterior margins are distinctly raised relative to the posterior part, which is composed of 3-5 nodes that are commonly higher than those and are inornate or bear a few poorly differentiated nodes. The tapered to the anterior, are partly fused, and extend to, or close to the pointed, and flat posterior platform is usually separated from the anterior part by or narrowly rounded to subrectangular posterior platform margin. marginal indentations. The free blade is about ⅓ of the total element The long free blade continues on the platform as an anterior carina of length and continues as a carina composed of 5-8 nodes that become 3-4 nodes that are lower than those located both to the anterior and larger towards the posterior end, where several may be partly fused. A posterior. The pit occupies a medial position beneath the platform, with generally narrow posterior platform brim is developed. The pit lies at a long keel extending posterior of it. or posterior of the center of the platform, and the keel extends beyond Comparisons: The posterior platform narrowing is much greater and it as a narrow rounded loop. more abrupt than in Parapetella columbiense, its likely precursor. The Comparisons: Compared with those of Parapetella riteri, the anterior overall shape of the elements is reminiscent of Acuminatella angusta, parapets are relatively longer and less differentiated, and the pit lies but that species has strongly nodose anterior margins. Elements farther to the posterior. Both species have variable anterior platform assigned to Quadralella spp. indet. have similar posterior narrowing, terminations, ranging from steep to sloped. but they are also anteriorly nodose, the posterior platform is much Stratigraphic occurrence: Common in the angusta-dylani Subzone shorter, and the pit is posterior in position. through middle subdivision of the parvus Subzone of the primitia Zone. Stratigraphic occurrence: Abundant in the angusta-dylani Subzone Parapetella cordillerense sp. nov. into middle subdivision of the parvus Subzone of the primitia Zone. Figure 53. 1-22 Parapetella destinae sp. nov. Derivation of name: Named for the Canadian Cordillera. Figure 54.10-27; 28-36 1973 [?] Epigondolella primitia Mosher - Mosher, p. 161, pl. 18, figs. Holotype: GSC 132858, Figure 53. 7-9. 3-5 (only). Type stratum: Bed 18f, within Pardonet Formation. 2012 “Metapolygnathus echinatus” sensu Orchard - Mazza et al. (b), p. 114, pl. 8, figs. 7a-c, 8a-c. Age: Middle subdivision of the parvus Subzone of the primitia Zone. Derivation of name: Named for my grand-daughter, Destiny. Diagnosis: Narrow, elongate P1 elements are characterized by distinctive partition into three parts of comparable length: an anterior Holotype: GSC 132875, Figure 54. 34-36. free blade, a relatively broad anterior platform, and a very narrow Type stratum: Bed 18h, within the Pardonet Formation. posterior platform. The biconvex anterior platform is twice as wide as the posterior part, being composed of raised platform parapets Age: Middle subdivision of the parvus Subzone of the primitia Zone. that are highest at or near their anterior end, where low nodes are Diagnosis: Small, very narrow, elongate P1 elements characterized by uncommonly developed. The slightly to strongly bowed posterior an anterior platform that bears short and high parapets on one or both platform is delineated anteriorly by abrupt narrowing beyond indented sides of the element at midlength; these may be apically sharp in early lateral margins. Narrow platform flanges flank the posterior carina, 77

Figure 53. 1-22. Parapetella cordillerense sp. nov. 1-3. GSC 132856, sample 18a. 4-6. GSC 132857, sample 18f. 7-9. GSC 132858, sample 18f. Holotype. 10-12. GSC 132859, sample 18e. 13-15. GSC 132860, sample 18c. 16-18. GSC 132861, sample 14a. 19-21. GSC 132862, sample 18c. 22. GSC 132863, sample 18d. Scale bar = 200 microns. growth stages but more typically round-topped and eventually buttress- johnpauli (q.v.). All these species are diminutive forms that were like in later growth stages. The straight to slightly curved platform to formerly united, but detailed comparison has led to their separation. the posterior of the parapets often narrows abruptly at indentations on Those assigned to P. destinae are differentiated on the basis of the pit both lateral margins, although in some elements the platform width is position into two morphotypes. maintained, and it may expand slightly towards the posterior margin, alpha morphotype which has a subrectangular outline. The free blade is between ⅓ and ½ Figure 54.10-27 of the total element length and descends onto the platform as 4-6 carinal nodes that maintain a constant height or only slightly rise towards Description: As for the species, with a pit that occupies a medial their posterior termination in front of a narrow platform brim. The pit or slightly anterior position beneath the platform. The free blade is occupies a medial to anterior position beneath the platform and within commonly ½ of the total element length. a broad keel that extends far to the posterior. Comparisons: The medial pit contrasts with the anterior pit of both Comparisons: These elements are similar to Parapetella cordillerense Parapetella destinae beta morphotype and P. johnpauli, which may in having a narrow posterior platform, but they have a more strongly also have longer blades. reduced anterior platform whereby the parapets are reduced to very Remarks: This morphotype includes the elements illustrated by Mazza short and high features. Primatella posteroglobosa has a similar shape et al. (2012b). but differs in its sharply defined anterior denticles. Elements assigned herein to Metapolygnathus parvus beta morphotype are distinguished Stratigraphic occurrence: Very common in the acuminata-prominens by lacking anterior parapets, although that species may have low nodes Subzone of the primitia Zone through basal quadrata Zone. This is one (compare Fig. 48. 41-43). of the few primitia Zone taxa that range into the quadrata Zone. Mazza et al. (2012b) give the range as Lower Lacian?, although they show it as Remarks: Orchard (2007c) combined diverse diminutive elements occurring only in the “boundary interval” (op. cit., fig. 2). in Metapolygnathus echinatus, and recognized them as potentially valuable in boundary definition as well as in trans-Panthalassa beta morphotype correlation. Mazza et al. (2012b) emphasized this in their recognition Figure 54. 28-36 of “M. echinatus” sensu Orchard at Pizzo Mondello. As discussed by Description: As for the species, with a pit that occupies a position Mazza et al. (2012b), the holotype of M. echinatus may be a juvenile beneath the junction of the platform and blade far anterior of the keel of M. communisti, and the present author concurs that the name should terminus. The free blade is about ½ of the total element length. The no longer be used for those elements that occur in the boundary interval parapets may be reduced and asymmetrically developed. of the parvus Subzone. However, a remarkably similar element to the holotype of M. echinatus was recovered from the Upper samueli Zone Comparisons: The beta morphotype differs from its probable of Haida Gwaii, a stratigraphic level that predates typical examples of forbearer, Parapetella destinae alpha morphotype, in its far anterior M. communisti (Orchard and Carter, 2013, fig. 3.10-13). pit position, longer blade, and sometimes reduced parapets. Similar P1 Other elements included in Metapolygnathus echinatus by Orchard elements with long blades and an anterior pit are P. johnpauli, which (2007c) are assigned in the present work to M. parvus and Parapetella has a laterally expanded posterior platform, and P. willifordi, which has 78

Figure 54. 1-3. Parapetella sp. nov. E. GSC 132864, sample 17b. 4-6. Parapetella sp. nov. F. GSC 132865, sample 18d. 7-9. Parapetella aff. destinae. GSC 132866, sample 14a. 10-27. Parapetella destinae sp. nov., alpha morphotype. 10-12. GSC 132867, sample10/02. 13-15. GSC 132868, sample 10/04. Holotype. 16-18. GSC 132869, sample 31. 19-21. GSC 132870, sample 29. 22-24. GSC 132871, sample 18a. 25-27. GSC 132872, sample 10/03b. 28-36. Parapetella destinae sp. nov., beta morphotype. 28-30. GSC 132873, sample 18h. 31-33. GSC 132874, sample 21d. 34-36. GSC 132875, sample 18h. Holotype. Scale bar = 200 microns. a pointed platform. Metapolygnathus parvus beta morphotype has a is equally broad at both anterior and posterior ends, with a distinctive similar platform outline but lacks anterior parapets. medial narrowing, and a roundly terminated posterior end. The anterior margins are distinctly raised as rounded, generally adenticulate Stratigraphic occurrence: Uncommon from the top of middle parapets. The free blade is about ⅓ the element length and continues as subdivision through upper subdivision of the parvus Subzone of the a carina composed of 5-7 nodes that become larger towards the posterior primitia Zone. end where several may be partly fused. A generally narrow posterior Parapetella aff. destinae platform brim is developed. The pit is in a medial position beneath the Figure 54. 7-9 platform, and the keel extends and broadens to the posterior. Remarks: A unique P1 platform element that features one adenticulate Comparisons: Parapetella columbiense, P. hillarae, and P. riteri parapet and a second that is divided into two nodes. This differs from differ in having a posterior platform that is narrower than the anterior the parapets of Parapetella destinae alpha morphotype, in which the platform, without posterior expansion as in the present species. The subrectangular posterior platform is also longer. Stratigraphic occurrence: Rare in the angusta-dylani Subzone of the Figure 55 (facing page). 1-9. Parapetella elegantula sp. nov. 1-3. primitia Zone. GSC 132876, sample 18d. 4-6. GSC 132877, sample 17c. 7-9. GSC 132878, sample 17c. Holotype. 10-27. Parapetella irwini sp. nov. 10- Parapetella elegantula sp. nov. 12. GSC 132879, sample 18f. 13-15. GSC 132880, sample 17c. 16- Figure 55. 1-9 18. GSC 132881, sample 18d. 19-21. GSC 132882, sample 18f. 22-24. Derivation of name: From elegan, -t (Latin) referring to the slender, GSC 132883, sample PHE-22. Pardonet Hill east. 25-27. GSC 132884, graceful shape of the platform. sample 18d. Holotype. 28-33. Parapetella pumilio sp. nov. 28-30. GSC 132885, sample 17c. 31-33. GSC 132886, sample 18c. Holotype. Holotype: GSC 132878, Figure 55. 7-9 34-36. Parapetella aff. pumilio. GSC 132887, sample 18d. 37-45. Type stratum: Bed 17c, within Pardonet Formation. Parapetella johnpauli sp. nov. 37-39. GSC 131168, sample PHE-23. Pardonet Hill east. Holotype. 40-42. GSC 132888, sample 18h. 43-45. Age: The acuminata-prominens Subzone of the primitia Zone. GSC 132889, sample 20b. 46-48. Parapetella sp. nov. D. 46-48. GSC Diagnosis: The P1 elements are characterized by a narrow platform that 132890, sample 20a. Scale bar = 200 microns. 79 80

Figure 56. 1-27. Parapetella hillarae sp. nov. 1-3. GSC 132891, sample 18. 4-6. GSC 132892, sample 14b. 7-9. GSC 132893, sample 18. 10- 12. GSC 132894, sample 17b. 13-15. GSC 132895, sample 18h. 16-18. GSC 132896, sample 13a. 19-21. GSC 132897, sample 18a. 22-24. GSC 132898, sample 14a. 25-27. GSC 132899, sample 14a. Holotype. Scale bar = 200 microns. hourglass shape of these elements resembles that seen in some platform. The unequal development of the two platform halves is Kraussodontus vancouverense, but those elements are broader and lack more pronounced than in the more elongate Parapetella columbiense. the well-differentiated anterior parapets. Elements of Quadralella pardoneti have a similar partition of the Stratigraphic occurrence: Uncommon in the angusta-dylani Subzone platform, but those elements have more quadrate posterior outlines, through the lower subdivision of the parvus Subzone of the primitia characteristically nodose anterior margins, and commonly a more Zone. posteriorly positioned pit. Parapetella hillarae sp. nov. Stratigraphic occurrence: Abundant in the angusta-dylani Subzone Figure 56. 1-27 through middle subdivision of the parvus Subzone of the primitia Zone. The species was also found in the Pelly Mountains of the Yukon 2006 Metapolygnathus n. sp. B - Orchard, pl. 7, figs. 1,1 12. Territory (Orchard, 2006), and in Nevada (Balini et al., 2014). 2014 Parapetella sp. nov. 1 - Balini et al., fig. 11.a1-3. Parapetella irwini sp. nov. Derivation of name: Named for Hillary Taylor in gratitude for her Figure 55. 10-27 long-standing assistance in conodont research. 2012 [p] Metapolygnathus communisti Hayashi - Mazza (b), p. 112, Holotype: GSC 132899, Figure 56. 25-27. 114, pl. 8, fig. 6 (only). Type stratum: Bed 14a, within Pardonet Formation. Derivation of name: Named for Steven Irwin, able field assistant early Age: The angusta-dylani Subzone of the primitia Zone. in this study, and later a supportive subdivision head. Diagnosis: P1 elements with a marked submedial platform constriction Holotype: GSC 132884, Figure 55. 25-27. separating two unequally developed halves. The anterior platform has Type stratum: Bed 18d, within Pardonet Formation. biconvex to subparallel margins that are moderately to strongly raised to form rounded to triangular parapets that may bear a few poorly Age: Lower subdivision of parvus Subzone of the primitia Zone. developed nodes. The posterior platform is consistently narrower than Diagnosis: P1 elements have elongate, subrectangular platforms with the anterior part and maintains its width through the rounded posterior subparallel lateral margins, a weak to moderate medial constriction, platform margin. The free blade is between ⅓ and ½ element length and a squared-off posterior margin. The anterior lateral margins rise and passes onto the platform as a carina that bears 4-6 nodes that tend gradually into high buttresses that become more pronounced with to become higher and more fused towards the posterior. The carina may growth, and often descend rapidly to meet the blade. The free blade extend very close to the posterior margin or terminate well in front of is about ⅓ total element length in mature specimens, and its denticles it, in which case a broad brim is developed. The pit is submedial in descend onto the platform as 4-7 carinal nodes that are at first low and position and clearly displaced anteriorly within the keel. then generally rise in height to the posterior, terminating a variable Comparisons: These elements are similar to Parapetella riteri alpha distance in front of the posterior margin; a small accessory node may morphotype but differ in their rounded rather than pointed posterior occur beyond. The pit is medial in position beneath the platform and has 81 a long posterior keel that is commonly bifurcated in adult specimens. angular or rounded postero-lateral corners. The width of the posterior Comparisons: The medial narrowing is reminiscent of that seen in platform is often narrower than, and does not exceed that of the anterior Parapetella elegantula, but the platform of the latter is posteriorly platform. The free blade is ~⅓ total element length and descends rounded rather than rectangular. Parapetella posterolata and P. sp. onto the platform as a row of 5-7 carinal nodes that are often highest nov. D have distinctly expanded posterior platforms that exceed the and partly fused at the posterior end of the carina; a narrow to wide breadth of the anterior platform, unlike the present species. P. irwini posterior platform brim is developed on which accessory carinal nodes resembles representatives of Metapolygnathus ex gr. communisti (q.v.) may occur. The pit underlies the middle of the platform and is followed but are distinguished by their high anterior parapets. Metapolygnathus by a broad keel that expands towards the posterior end where it might angustus Kozur has a similar platform shape and also the suggestion of be bifurcated in larger specimens. a low anterior parapet, but its pit is far forward of medial in position. Comparisons: These elements have a much stronger medial Stratigraphic occurrence: Uncommon in the acuminata-prominens constriction than other species with an hourglass shaped platform, e.g., Subzone through basal middle subdivision of the parvus Subzone of Parapetella elegantula and Kraussodontus vancouverense. They differ the primitia Zone. Mazza et al. (2012b) illustrated a specimen here from the much broader Parapetella posterolata and P. sp. nov. D, which assigned to this species in sample NA37 at Pizzo Mondello, which is have expanded posterior platforms that exceed the width of the anterior from within their boundary Zone. platform. Parapetella johnpauli sp. nov. Remarks: Orchard (2007c) combined these elements with larger Figure 55. 37-45 specimens of Quadralella praecommunisti curvata in the belief that the former represented early growth stages of the latter. However, although 2001 [p] Metapolygnathus echinatus (Hayashi) - Orchard (c), pl. 1, they have a similar occurrence, they are separated here because the figs. 7-9 (only). characteristic posterior expansion of the present species persists in Derivation of name: Named for John-Paul Zonneveld for his lead in larger specimens. field logistics and stratigraphy of Black Bear Ridge. Stratigraphic occurrence: Abundant in the angusta-dylani Subzone Holotype: GSC 132888, Figure 55. 37-39. through middle subdivision of the parvus Subzone of the primitia Zone. Type stratum: Bed 18h, within Pardonet Formation. Parapetella posterolata sp. nov. Figure 57. 31-39 Age: Middle subdivision of parvus Subzone of the primitia Zone. Derivation of name: From postero-, lat (Latin): refers to wide posterior Diagnosis: Diminutive P1 elements in which the platform is less than platform. ½ the total element length. The platform is broad at both its anterior and, particularly, posterior ends, and narrows medially where a strong Holotype: GSC 132909, Figure 57. 31-33. constriction occurs. The anterior margins are raised as short parapets Type stratum: Bed 18d, within Pardonet Formation. that may appear as rounded nodes; additional nodes may occur on the outer margin. The posterior platform has a triangular outline with Age: Lower subdivision of the parvus Subzone of the primitia Zone. extended postero-lateral corners and commonly an indentation in the Diagnosis: The P1 elements are characterized by a large platform with posterior margin. The long free blade extends onto the platform as a medial constriction that separates an anterior part from a strongly 2-5 carinal nodes that terminate near the posterior margin; subdued expanded posterior platform, the maximum breadth of which, at the secondary carinal ridges may be developed. On the lower surface, the posterior margin, exceeds the width of the anterior platform by up to pit is at or very close to the anterior platform margin-blade junction, 50%. The anterior platform margins bear parapets that have an upwardly and a bifid keel extends posteriorly beyond it. convex profile and may bear a few incipient to weakly differentiated Comparisons: These elements have a similar platform outline to nodes. To the posterior, a constriction reduces the platform width Parapetella lanei but are smaller, have a longer blade, and an anterior slightly, but almost immediately the lateral margins expand to define a pit. The pit position is like that seen in other diminutive species that broad subtriangular posterior platform with strongly extended postero- differ in platform outline, e.g., the subrectangular Parapetella destinae lateral corners. The free blade is ~⅓ total element length and descends beta morphotype, and the pointed P. willifordi. onto the platform as a row of 4-6 carinal nodes that are often highest and partly fused at the posterior end of the carina; a wide posterior Remarks: This species appears to be a derivative of the larger platform brim is developed. The pit underlies the middle of the platform Parapetella lanei through a general diminution in size, and particularly or slightly posterior of it and is followed by a broad keel that expands through reduction of the anterior platform, thereby producing both a towards the posterior end where it bifurcates and extends to each relatively longer free blade and a more anteriorly located pit. Most postero-lateral corner. specimens retain the overall outline of the older species, albeit in a miniature form. Comparisons: These elements are broader than Parapetella lanei and have a posterior platform with a breadth that, in contrast to the latter, Stratigraphic occurrence: Very common in the middle and upper exceeds that of the anterior part. P. sp. nov. D is similar in outline but subdivisions of the parvus Subzone of the primitia Zone. has more differentiated parapets and a more anteriorly positioned pit. Parapetella lanei sp. nov. P. johnpauli has a similar shape but is much smaller with a longer free Figure 57. 1-30 blade and a far anterior pit. 2007 [p] Metapolygnathus n. sp. P - Orchard (c), pl. 2, figs. 34-36 Remarks: It is possible that some smaller specimens of Parapetella (only). lanei are earlier growth stages of this species, although both species 2010 Metapolygnathus n. sp. P - Orchard (a), fig. 13. 17. occur as comparably large elements. Stratigraphic occurrence: Very common in the acuminata-prominens Derivation of name: Named for Larry Lane, GSC project manager Subzone through the lower subdivision of the parvus Subzone of the under which this study was completed. primitia Zone. Holotype: GSC 132907, Figure 57. 25-27. Parapetella prominens Orchard Type stratum: Bed 18f, within Pardonet Formation. Figure 58. 1-35 Age: Middle subdivision of parvus Subzone of the primitia Zone. 2013 Parapetella prominens sp. nov. - Orchard, p. 452, fig. 3.25-

Diagnosis: The P1 elements are characterized by a platform with a 3.27. strong medial constriction separating a broad anterior part from an Diagnosis: The P1 elements of this species have a platform that is about expanded posterior platform, producing an hourglass outline. The twice as long as broad, and a free blade that is between ⅓ and ½ of anterior platform has strongly elevated marginal parapets with a the total element length. The long axes of the elements are straight to generally rounded profile that may be interrupted by the presence of curved, and the posterior platform outlines include rectangular, rounded, weak nodes. The strong medial constriction reduces the platform width and tapered elements. All specimens have high anterior parapets of up to ½, and to the posterior it expands to a subtriangular shape with variable shape that double the height of the platform and are clearly 82 83

Figure 58. 1-12. Parapetella prominens circulare subsp. nov. 1-3. GSC 132913, sample 18f. 4-6. GSC 132914, sample 18h. 7-9. GSC 132915, sample 17c. 10-12. GSC 132916, sample 17b. Holotype. 13-24. Parapetella prominens prominens Orchard. 13-15. GSC 132917, sample 18a. 16-18. GSC 132918, sample 10/03. 19-21. GSC 132919, sample 17b. 22-24. GSC 132606, sample 18e. Holotype. 25-33, ?34, 35. Parapetella prominens angulare subsp. nov. 25-27. GSC 132920, sample 18e. Holotype. 28-30. GSC 132921, sample 14a. 31-33. GSC 132922, sample 18c. 34, 35. GSC 132923, sample 18c. Scale bar = 200 microns.

differentiated from the relatively flat posterior platforms. The carina is Figure 57 (facing page). 1-30. Parapetella lanei sp. nov. 1-3. composed of 4-6 nodes that are commonly highest at the posterior end GSC 132900, sample 18f. 4-6. GSC 132901, sample 17c. 7-9. GSC where they may be partly fused; platform brims are of variable width, 131174, sample 13a. 10-12. GSC 132902, sample 18f. 13-15. GSC and may include an accessory denticle. The pit underlies the center of 132903, sample 18d. 16-18. GSC 132904, sample 18f. 19-21. GSC the platform, far anterior of the posterior keel termination that reflects 132905, sample 13c. 22-24. GSC 132906, sample 18h. 25-27. GSC the shape of the platform (Orchard, 2013). 132907, sample 18f. Holotype. 28-30. GSC 132908, sample 18f. 31- Comparisons: The very prominent parapet separates this species from 39. Parapetella posterolata sp. nov. 31-33. GSC 132909, sample others of similar outline. 18d. Holotype. 34-36. GSC 132910, sample 18d. 37-39. GSC 132911, sample 17c. 40-42. Parapetella sp. nov. G. GSC 132912, sample 31. Remarks: This species is divided into three subspecies based on the Scale bar = 200 microns. differing posterior platform outlines. 84 Parapetella prominens prominens subsp. nov. Figure 55. 28-33 Figure 58. 13-24 Derivation of name: From pumil, -io (Latin), dwarf, referring to the 2013 Parapetella prominens sp. nov. - Orchard, p. 452, fig. 3.25- diminutive nature of the species. 3.27. Holotype: GSC 132886, Figure 55. 31-33. Derivation of name: As for species. Type stratum: Bed 18c, within the Pardonet Formation. Holotype: As for species. GSC 132606, Figure 58. 22-24. Age: Lower subdivision of the parvus Subzone of the primitia Zone. Type stratum: Bed 18e, within the Pardonet Formation. Diagnosis: Diminutive P1 elements with strongly reduced, Age: Lower subdivision of parvus Subzone of the primitia Zone. asymmetrical platforms. The anterior platform occurs as lateral flanges Diagnosis: P1 elements of this subspecies have a sub-rectangular on which tiny raised geniculation points may or may not occur, and a platform and a straight posterior margin that may be either perpendicular less reduced, asymmetric posterior platform that is the broadest part of or oblique in relation to the posterior carina. The long axis of the the element. Separating the two, the platform is almost entirely reduced elements is straight to curved, and the lateral margins are sub-parallel. by a medial constriction. The free blade is ~⅓ total element length, and its denticles merge into the carinal nodes, which remain high before Comparisons: These elements are similar in shape to Parapetella terminating in front of the posterior margin. The pit is medial, lying beattyi but they have a far more differentiated parapet in adult specimens. beneath the strongly constricted part of the platform. Some Quadralella species also have a similar outline, but they lack a parapet entirely, as does the superficially similar Metapolygnathus Comparisons: These elements differ from their likely precursor dylani. Parapetella lanei in being much narrower and lacking a well-developed anterior platform as well as a reduced posterior platform. Whereas Stratigraphic occurrence: Uncommon in the acuminata-prominens Parapetella johnpauli retains the platform shape of the former species Subzone through the asymmetrica-Norigondolella sp. Subzone of the but developed an anterior pit, the derivation of the present species primitia Zone. through lateral reduction preserved a medial pit. Parapetella prominens angulare subsp. nov. Stratigraphic occurrence: Rare in the acuminata-prominens Subzone Figure 58. 25-33, ?34, 35 through lower subdivision of the parvus Subzone of the primitia Zone. Holotype: GSC 132920, Figure 58. 25-27. Parapetella aff. pumilio Derivation of name: From anguli- (Latin) angled, referring to the Figure 55. 34-36 offset posterior platform. Description: A unique P1 element with a long high blade and a tiny vestigial platform developed in the posterior ¼- of the element. The Type stratum: Bed 18e, within the Pardonet Formation. 1/5 apparently very large but broken cusp is the penultimate denticle and Age: Lower subdivision of the parvus Subzone of the primitia Zone. lies above a pit that lies within a broad furrowed keel with a posterior Diagnosis: As for the species but characterized by a platform that tapers rounded loop. from about its midlength to a narrowly pointed to rounded posterior Comparisons: This element differs from Parapetella pumilio, a likely margin that is offset to the inner side. The lateral margins are plano- precursor, in lacking any trace of an anterior platform. The high blade, convex or concavo-convex as a result of the long axis being curved. large cusp, and character of the pit and loop resemble that of a “naked” Comparisons: These elements differ from Parapetella clareae in gondolellid. Unlike Metapolygnathus parvus gamma morphotype, this their stronger parapet and absence of posterior-lateral expansion of the species has a posterior platform and different denticulation. outer platform. Kraussodontus urbanae is much longer and has noded Stratigraphic occurrence: Rare in the lower subdivision of the parvus anterior margins rather than well differentiated parapets. Subzone of the primitia Zone. Remarks: A unique specimen (Fig. 58. 34, 35) is tentatively regarded Parapetella riteri sp. nov. as an extreme example of this subspecies. It differs in having a more Figure 59. 4-26; 27-35 strongly incurved posterior “tongue” and a high carina that extends to the pointed posterior tip of the platform. Derivation of name: Named for the late Frank Riter of Hudson Hope, who provided water transport on Williston Lake during the early stages Stratigraphic occurrence: Uncommon in the acuminata-prominens of this study. Subzone through lower subdivision of the parvus Subzone of the primitia Zone. Holotype: GSC 131178, Figure 59. 21-23. Parapetella prominens circulare subsp. nov. Type stratum: Bed 14a, within the Pardonet Formation. Figure 58. 1-12 Age: The angusta-dylani Subzone of the primitia Zone. Holotype: GSC 132916, Figure 58. 10-12. Diagnosis: The P1 elements have a variable platform length-to-breadth Derivation of name: From circul- (Latin), referring to the rounded ratio, between 1:1 and 2.5:1, are broadest at their anterior end and posterior platform margin. tapered progressively to a narrowly rounded posterior tip. The lateral margins are mostly straight, but marginal indentation may occur on Type stratum: Bed 17b, within the Pardonet Formation. the posterior margin of the upstanding anterior parapets, which may Age: The acuminata-prominens Subzone of the primitia Zone. be symmetrical and rounded in profile, or asymmetrical with a steep anterior edge. The anterior parapets may have incipient node formation Diagnosis: As for the species, but the P1 elements are characterized but are generally inornate. The free blade is between ⅓ and ½ element by a commonly ovoid platform with a straight long axis, and a well- length and passes posteriorly into 2-6 carinal nodes, the posteriormost rounded posterior outline. The platform elements are commonly of which (or several adjacent, partly fused nodes) is the largest. The pit biconvex with a maximum width at mid-length, or less commonly they lies close to the center of the platform and has a pointed posterior keel have sub-parallel margins. extending behind it. Comparisons: The rounded platform shape is like that of Kraussodontus Comparisons: The overall platform shape of these P1 elements roberti beta morphotype, but those elements have a relatively flat is like that of Acuminatella spp., but the latter differ in having platform and lack strongly differentiated parapets. distinct anterior nodes and a posterior carina. The species resembles Stratigraphic occurrence: Uncommon in the acuminata-prominens Parapetella columbiense but the present species is shorter, and has Subzone through middle subdivision of the parvus Subzone of the more differentiated parapets. The elements differ from P. hillarae in primitia Zone. their tapered and pointed posterior platforms. Parapetella pumilio sp. nov. Remarks: Two morphotypes are differentiated: 85

Figure 59. 1-3. Parapetella aff. riteri. 1-3. GSC 132924, sample 8. 4-23, ?24-26. Parapetella riteri sp. nov., alpha morphotype. 4-6. GSC 132925, sample 14b. 7-9. GSC 132926, sample 17b. 10-12. GSC 132927, sample 18a. 13, 14. GSC 132928, sample 17b. 15-17. GSC 132929, sample 18a. 18-20. GSC 132930, sample 18a. 21-23. GSC 131178, sample 15. Holotype. 24-26. GSC 131177, sample 14a. 27-35. Parapetella riteri sp. nov., beta morphotype. 27-29. GSC 131175, sample 16. 30-32. GSC 132931, sample 14a. Holotype. 33-35. GSC 132932, sample 13a. Scale bar = 200 microns. alpha morphotype into basal middle subdivision of the parvus Subzone of the primitia Figure 59. 4-23, ?24-26 Zone. 2007 Metapolygnathus n. sp. K - Orchard (c), pl. 2, figs. 31-33, beta morphotype ?40-42. Figure 59. 27-35 Description: This morphotype has an elongate pointed platform, 2007 [p] Metapolygnathus n. sp. Y - Orchard (c), pl. 2, figs. 28-30 commonly with a length-to-breadth ratio of 2:1, and a high triangular (only). parapet on each anterior margin in which the apex is at the subvertical 2010 Metapolygnathus n. sp. Y - Orchard (a), fig. 13. 21. anterior platform margin. The free blade is between ⅓ and ½ element length and passes into 3-4 variously spaced carinal nodes. Description: This morphotype has a relatively long, roundly pointed platform with a length-to-breadth ratio of 2.5:1, and high parapets with Remarks: A large specimen (Fig. 59. 24-26) is questionably included a symmetrical rounded profile and a central high-point. The blade is here because, although it has identical parapets, it has a longer, less about ⅓ element length and passes into 5-6 partly fused, carinal nodes. pointed posterior platform, and seven carinal nodes. It is tentatively regarded as a late growth stage of this species. Stratigraphic occurrence: Abundant in the angusta-dylani Subzone through the acuminata-prominens Subzone of the primitia Zone. Stratigraphic occurrence: Abundant in the angusta-dylani Subzone 86 Parapetella aff. riteri Diagnosis: P1 elements with very short platforms, broadest in the Figure 59. 1-3 anterior or medial parts, and with a length-to-breadth ratio close to 1:1. Description: This unique specimen corresponds to the beta morphotype The anterior parapets are subtriangular with a sloping anterior edge, of P. riteri except that the outer anterior margin has 3 low nodes rather a high apex, and a less steeply sloping posterior profile. The blade is than a high parapet with a subtriangular profile, as is developed on the about ½ total element length and passes onto the platform as 2-3 isolated inner margin. The pit lies posterior of the platform midlength close to nodes separated from the posterior margin by a brim. The amplitude of the posterior end of the keel. the parapets and the width of the brim increase with growth in these elements. The pit lies a little anterior of the platform midlength. Remarks: This element differs from typical elements of Parapetella riteri in both its unequally developed anterior margins and more Comparisons: These elements appear related to Parapetella riteri posterior pit position. The single parapet is well differentiated but has but are much shorter and relatively broad. In profile, the parapets are neither a rounded profile like that of the alpha morphotype, nor a steep pointed but their anterior edge is less steep than in the beta morphotype anterior edge like that of the beta morphotype. Its stratigraphic level of P. riteri. P. willifordi differs in its relatively longer free blade and its suggests it is a precursor to typical P. riteri. more anteriorly located pit. Stratigraphic occurrence: Rare in the sagittale-beattyi Subzone of the Stratigraphic occurrence: Uncommon in the angusta-dylani Subzone primitia Zone. of the primitia Zone. Parapetella rubae sp. nov. Parapetella willifordi sp. nov. Figure 60. 1-9 Figure 60. 10-27 Derivation of name: Named for Ruby Merriman in gratitude for her Derivation of name: Named for Kenneth Williford, who undertook support during the course of this study. chemostratigraphic studies at Black Bear Ridge. Holotype: GSC 132934, Figure 60. 4-6. Holotype: GSC 132939, Figure 60. 19-21. Type stratum: Bed 14b, within Pardonet Formation. Type stratum: Bed 18h, within Pardonet Formation. Age: The angusta-dylani Subzone of the primitia Zone. Age: Middle subdivision of the parvus Subzone of the primitia Zone. Diagnosis: These generally diminutive P1 elements have short, subtriangular to oval shaped platforms with a length-to-breadth ratio

Figure 60. 1-9. Parapetella rubae sp. nov. 1-3. GSC 132933, sample 14b. 4-6. GSC 132934, sample 14b. Holotype. 7-9. GSC 132935, sample 14b. 10-27. Parapetella willifordi sp. nov. 10-12. GSC 132936, sample 21. 13-15. GSC 132937, sample 18h. 16-18. GSC 132938, sample 20f. 19-21. GSC 132939, sample 18h. Holotype. 22-24. GSC 132940, sample 18c. 25-27. GSC 132941, sample 21d. Scale bar = 200 microns. 87 lying between 1:1 and 1.5:1, and have long blades equal to ½ or 2/3 Stratigraphic occurrence: Rare in the acuminata-prominens Subzone of the total element length. The platforms are tapered to a sharp or of the primitia Zone. narrowly rounded posterior margin, and the lateral platform margins Parapetella sp. nov. D are straight to biconvex; commonly, the inner platform is broader with Figure 55. 46-48 a more convex margin. The anterior lateral margins are raised and produced into parapets of varying height; exceptionally the parapet is Description: A P1 element with a very short, subtriangular platform, node-like. The long free blade passes into a carina composed of 2-4 secondary posterior carinae, a blunt anterior node on each margin, and nodes that almost reach the posterior tip in small specimens but stop an anterior pit at the platform-blade junction. The free blade is missing well in front of it in larger elements. The pit is at, or close to the junction but was probably long. of the platform and blade, and a pointed keel extends under most of the Remarks: This element resembles the posterior platform of Parapetella platform. posterolata and the similar but diminutive P. johnpauli, but it lacks an Comparisons: The P1 platform shape is similar to that of Parapetella anterior platform. It may have been derived from those species through rubae, but both the blade length and pit position differ. The generally complete reduction of the anterior platform. pointed platform is reminiscent of P. riteri, which is a probable Stratigraphic occurrence: Uncommon in the middle and upper precursor. The platform of the diminutive Metapolygnathus parvus is subdivisions of the parvus Subzone of the primitia Zone. similar, but pointed forms are excluded from that species. Parapetella sp. nov. E Stratigraphic occurrence: Abundant in the middle and upper Figure 54. 1-3 subdivisions of the parvus Subzone of the primitia Zone. Remarks: This unique specimen differs from Parapetella cordillerense Parapetella sp. nov. A in bearing a very short anterior platform that is less than ½ the length Figure 51. 19-27 of the narrow posterior platform. It is also close to P. destinae, but the Description: The P1 elements are short, asymmetrically ovoid, with a posterior carina is longer and posterior platform relatively narrower. length-to-breadth ratio of 2:1, plano-convex lateral platform outlines, Stratigraphic occurrence: Rare in the acuminata-prominens Subzone and a rounded posterior margin. The anterior lateral margins gradually of the primitia Zone. rise to high geniculation points at the anterior end of inornate parapets. The blade is equal to about ½ element length, and the carina is composed Parapetella sp. nov. F of 3-5 nodes that terminate in a large node that becomes increasingly Figure 54. 4-6 distant from the posterior margin as the posterior brim lengthens. The Remarks: This unique element is similar to Parapetella lanei but pit is submedial under the platform, and anteriorly shifted in the keel, has an anterior platform that is laterally reduced on each margin to a which has a pointed to squared-off termination. low narrow flange lacking a distinctive parapet. The narrow posterior Comparisons: These elements are similar to Kraussodontus roberti platform is slightly expanded, and the carina is very high to the posterior beta morphotype but differ in having a curved asymmetric platform with of the subcentral pit. more prominent anterior parapets, similar to those seen in Parapetella Stratigraphic occurrence: Rare in the acuminata-prominens Subzone beattyi. of the primitia Zone. Remarks: These abbreviated elements are similar to, and partly co- Parapetella sp. nov. G occur with, Parapetella beattyi, P. broatchi, and P. clareae, which Figure 57. 40-42 collectively mimic shape variation in those elements here described as subspecies of P. prominens. The former are named as individual species Description: A unique P1 element characterized by a large platform because the shape variation is more extreme and they appear to have with a medial constriction that separates an anterior part from a strongly differing ranges. expanded posterior platform that exceeds the width of the anterior. The anterior platform is less than ½ the total platform length and consists Stratigraphic occurrence: Common in the sagittale-beattyi Subzone of strongly differentiated and high parapets that bear incipient nodes. through the angusta-dylani Subzone of the primitia Zone. To the posterior, a constriction reduces the platform width and then the Parapetella sp. nov. B lateral margins expand progressively towards extended postero-lateral Figure 51. 28-32 corners that mark the maximum platform width. The free blade is ~⅓ total element length and descends onto the platform as a row of five Description: Small, squat P1 elements with a length-to-breadth ratio carinal nodes that end in front of a wide posterior platform brim. The approaching 1:1. The platform has a broad anterior half with high, pit underlies the anterior half of the platform and is followed by a broad upturned lateral margins forming parapets, posterior to which a medial keel that expands towards the posterior end where it bifurcates and constriction separates a short tapered posterior platform that is about ½ extends towards each postero-lateral corner. the width of the anterior; a large cusp lies at its centre. The free blade is about ½ the length of the element. The pit is terminal in the keel and Comparisons: This species closely resembles Parapetella posterolata, positioned beneath the platform constriction. from which it probably developed, but differs from it in having more pronounced anterior parapets that are higher, shorter, and have more Comparisons: Similar small P1 elements of Quadralella? stephanae precipitous margins, and a more anteriorly located pit. have nodose anterior margins rather than parapets. See also Parapetella sp. nov. C. Stratigraphic occurrence: Rare near the top of the asymmetrica- Norigondolella sp. Subzone of the primitia Zone, immediately prior to Stratigraphic occurrence: Uncommon in the sagittale-beattyi the FAD of E. quadrata. Subzone into basal angusta-dylani Subzone of the primitia Zone. Genus Primatella Orchard Parapetella sp. nov. C Figures 21-25; 61-74 Figure 51. 33-38 2013 Primatella gen. nov. - Orchard, p. 452-3. Description: Small, squat P1 elements with a length-to-breadth ratio of 1-1.5:1. The platform has a broad anterior half with high, well Type species: Epigondolella primitia Mosher, 1970 differentiated parapets. A strong medial constriction separates a lobate Diagnosis: Anteriorly ornate, posteriorly downarched P1 platform posterior platform that is about ½-2/3 of the width of the anterior. The elements of variable outline with generally inornate posteriors and free blade is about ½ the length of the element and passes into 3-4 moderately high, apically blunt to sharp anterior marginal denticles that carinal nodes, the largest of which is terminal at the beginning of, are commonly equal to between 20% and 40% of the total depth of or within, the posterior platform. The pit is medial in position and the element measured from the denticle tip to the basal keel margin. subterminal within the keel. Dense microreticulation covers both the platform and round-topped Comparisons: These elements are similar to those of Parapetella sp. nodes. The carina usually terminates in front of the posterior margin, nov. B, but they have a broader, more lobate posterior platform, more and a platform brim is generally developed. The pit position is variable differentiated parapets, and a more anteriorly located pit. but commonly medial or posterior, and only rarely anterior (Orchard, 2013). 88 Description: The P1 platform elements are generally long with a respects, so the new genus Primatella was introduced (Orchard, 2013). length-to-breadth ratio ranging from 1.5:1 to 3:1. In some species, Rare Primatella with relatively long platforms and posterior pits the elements are subrectangular in outline, but others show a distinct first appear with the last Carnepigondolella, which are characterized, medial constriction, particularly in early growth stages. The posterior in contrast, by short, abbreviated platforms and medial pits. Although margin, although often straight, may be obliquely truncated, rounded, both genera are ornate, the latter is not regarded as ancestral to or postero-laterally expanded. In lateral view, Primatella species are Primatella, which is thought to have developed from Quadralella arched with a downturned posterior platform. through increasing differentiation of anterior nodes that are relatively The platform is often divided into two subequal parts, which is large compared with those found in typical Carnepigondolella. The particularly evident in those elements with a medial constriction. The first species of Primatella has a posterior pit, and round-topped and anterior part bears 2-6, commonly three, moderately sized marginal poorly differentiated anterior nodes. During the initial diversification nodes or denticles on each margin; the two margins may be unequally of the stock, both short and long subquadrate forms, and others with developed. The denticles are usually well differentiated and have either a platform constriction, can be differentiated and seemingly give rise sharp apices or may be round-topped, particularly in stratigraphically to similarly shaped successor species with more pronounced anterior older species and in large specimens of descendant species. The denticles and a more anteriorly located platform constriction and pit. posterior platform is generally unornamented, but exceptionally may Carinal characteristics and length are variable, and serve to distinguish bear scalloped or incipiently nodose margins. Compact microreticulae several species introduced here to help clarify the correlation potential cover the platform margins and nodes. of these various configurations. Twenty-two species ofPrimatella , plus The free blade is approximately 1/3 total unit length with closely ten morphotypes are described below. spaced, partly fused denticles forming an arcuate upper profile, with its maximum height at midlength or, more commonly, near its anterior end. Stratigraphic occurrence: The range of Primatella broadly The denticles descend onto the platform and continue as often small corresponds to that of the primitia Zone at Black Bear Ridge. They nodes that increase in size to the posterior. The carina, consisting of are relatively uncommon in the sagittale-beattyi Subzone and then 3-6 nodes, is variably developed to the posterior of the ornate anterior become overwhelmingly dominant in the asymmetrica-Norigondolella platform and forms the basis for speciation. In many specimens the sp. Subzone of the primitia Zone. posteriormost carinal node is the largest, although it is not necessarily Primatella asymmetrica Orchard the cusp. In some younger species, the terminal carinal node lies close Figure 61. 4-27 to the centre of the platform; in others the carina extends close to the posterior margin. A posterior platform brim varies from wide to very 2013 Primatella asymmetrica sp. nov. - Orchard, p. 454-55, fig. 4. narrow. 19-21. The pit varies in position from the posterior of midlength and 2013 Primatella asymmetrica Orchard - Carter and Orchard, fig. 7. subterminal within the keel in early forms, to subcentral and anteriorly 16-18. shifted within the keel in younger forms. In general, the pit lies beneath 2014 Primatella asymmetrica Orchard - Balini et al., fig. 11.k1-3, the platform constriction in those elements in which it is developed. l1-3. Exceptionally, the pit lies to the anterior of the platform centre. Holotype: GSC 132616, Figure 61. 25-27 Comparisons: The ornate character of Primatella elements distinguishes them from the inornate or weakly ornate representatives of Type stratum: Bed 18h, from within the Pardonet Formation. Quadralella, Kraussodontus, Metapolygnathus, and Parapetella. The Age: Middle subdivision of the parvus Subzone Subzone of the primitia P1 elements of Acuminatella differ in being narrower, having smaller Zone. and more numerous platform nodes, and typically a long and high carina that extends to the pointed posterior platform margin. Epigondolella Diagnosis: A distinctive P1 element with an expanded, strongly convex has larger and sharper anterior denticles, a lower lateral profile that is outer platform margin that meets the straight inner platform margin at generally stepped up posteriorly rather than downturned, a pit that is one postero-lateral corner: this imparts an asymmetric lobate posterior consistently medial in position, and less compact microreticulation on outline. Rarely, the posterior margin is crenulated but otherwise it is the platform. Carnepigondolella species have smaller rounded nodes unornamented. The anterior platform margins bear 3-5 discrete sharp or sharp denticles at their less steep anterior platform margins, and denticles or, in larger specimens, round-topped discrete nodes on each commonly also on their posterior platform. margin. The blade is about ⅓ element length and continues as a carina composed of 4-5 discrete nodes that either terminate well in front of the Remarks: Early studies of the Pardonet Formation conodonts posterior margin and a broad platform brim, or as an additional isolated occurring in Williston Lake outcrops led to the characterization of posterior node in front of a narrow brim. The pit is subcentral and lies an “Epigondolella primitia population” by Orchard (1983, fig. 2). within an arcuate keel that extends posteriorly far beyond it (Orchard, This composite fauna included diverse morphotypes that commonly 2013). co-occur in the abundant faunas of the latest Carnian and earliest Norian age in that area. Initially, Kozur (1989a, p. 402, 403) regarded Comparisons: The asymmetrical and lobate posterior outline and lack “Epigondolella primitia” as an example of E. abneptis (Huckriede), but of a posterior carina distinguishes this species from others assigned to re-illustration of the holotype by Orchard (1991b) showed it had a more Primatella. Some elements referred to Acuminatella acuminata have anteriorly located pit, less developed anterior denticles, a distinctive a slightly asymmetric but much narrower posterior platform and a platform microreticulation, and a differing lower surface profile. The carina that continues to the posterior tip. Parapetella clareae has a very anterior location of the pit in particular led Orchard (1991b) to refer the similar shape but lacks anterior ornament. species to Metapolygnathus because such a feature is characteristic of Stratigraphic occurrence: Common from the base of the parvus the type species of that genus, M. communisti Hayashi. Subzone through the asymmetrica-Norigondolella sp. Subzone of the Among the “Epigondolella primitia population” of Orchard (1983), primitia Zone. The species is also known in both Haida Gwaii (Carter several independent species have subsequently been differentiated. and Orchard, 2013) and Nevada (Balini et al., 2014). Kozur (2003) introduced “Epigondolella” orchardi for medially constricted P1 elements with the terminal node of the carina in a Primatella aff. asymmetrica Orchard central position and a more posteriorly situated pit; “Metapolygnathus” Figure 61. 1-3 mersinensis was introduced by Kozur and Moix (in Moix et al., 2007) 2012 Metapolygnathus mersinensis Kozur and Moix - Mazza et on the basis of its shorter platform and differing posterior carina. In al.(b) , p. 116-17, pl. 4, fig. 4. this work, additional species are introduced for elements with differing posterior carinas and different platform shapes. The stratigraphically Remarks: These uncommon P1 elements have an asymmetrical oldest species have a more posteriorly located pit and round-topped posterior platform outline, similar to but less pronounced than that of and/ or less differentiated nodes rather than sharp anterior denticles. Primatella asymmetrica. In common with the latter, the carina terminates Primatella primitia sensu stricto is restricted to forms with three well in front of the posterior margin and the pit-keel configuration closely spaced posterior carinal nodes. All these species are united is the same. In contrast, the anterior platform nodes of the present in having well developed anterior platform nodes or denticles, and species are less differentiated and the blade is shorter. Its stratigraphic strong microreticulation over both platform and nodes. However, position suggests it may be the precursor of P. asymmetrica. The carina they correspond neither to Metapolygnathus nor Epigondolella in key 89

Figure 61. 1-3. Primatella aff. asymmetrica Orchard. GSC 132942, sample 4. 4-27. Primatella asymmetrica Orchard. 4-6. GSC 132943, sample 20. 7-9. GSC 132944, sample 21h. 10-12. GSC 132945, sample 24. 13-15. GSC 132946, sample 18h. 16-18. GSC 132947, sample PHE- 24. Pardonet Hill east. 19-21. GSC 132948, sample 30. 22-24. GSC 132949, sample 21h. 25-27. GSC 132616, sample 18h. Holotype. 28-30. Primatella sp. nov. A. 28-30. GSC 132950, sample 31. Scale bar = 200 microns. configuration suggests a possible relationship also toP. ovale and P. ex 114, 116, pl. 8, fig. 11. gr. pseudoechinata. 2014 Primatella sp. nov. 6 - Balini et al., fig. 10 j1. Stratigraphic occurrence: Rare in the spenceri Subzone of the Derivation of name: Refers to the bifid nature of the posterior margin samueli Zone through the angusta-dylani Subzone of the primitia Zone. of the P1 element. A similar specimen from Pizzo Mondello comes from sample FNP53 Holotype: GSC 132956, Figure 62. 13-15. (Mazza et al., 2012b), apparently within the samueli Zone. Type stratum: Bed 07-22, within the Pardonet Formation. Primatella bifida sp. nov. Figure 62. 1-15 Age: The asymmetrica-Norigondolella sp. Subzone of the primitia Zone. 2012 [?] Metapolygnathus linguiformis Hayashi - Mazza et al. (b), p. 90

Figure 62. 1-15. Primatella bifida sp. nov. 1, 2. GSC 132951, sample 23. 3-5. GSC 132952, sample 18d. 6-8. GSC 132953, sample 18d. 9. GSC 132954, sample 23. 10-12. GSC 132955, sample 21g. 13-15. GSC 132956, sample 07-22. Holotype. Scale bar = 200 microns. Diagnosis: The P1 elements have broad rectangular to subtriangular Metapolygnathus linguiformis Hayashi, as interpreted by Mazza et platforms characterized by distinctive outgrowths of the postero-lateral al. (2012b), is also characterized by a bifid posterior margin. Although corners of the platform that are accentuated by a medial sinus on the the holotype of that species is badly damaged, it evidently lacks anterior posterior margin. Smaller growth stages may have a medial platform marginal nodes. The specimen illustrated by Mazza (op. cit.) does have constriction, which is less evident with later growth. The anterior a few low nodes, and is therefore questionably included in synonymy margin has strong and discrete, pointed to apically rounded denticles here. Both “Neogondolella” bisecta Igo 1989 and “Metapolygnathus” and nodes. There are 1-2 well separated carinal nodes posterior of slovenicus Ramovs 1994 are similar to the holotype of M. linguiformis the cusp, and occasionally a suggestion of a secondary carina. A wide in generally lacking anterior nodes: these three taxa may be synonyms. posterior platform brim is commonly developed. The pit is medial, and Stratigraphic occurrence: Common in the parvus Subzone through the posterior keel bifurcates about mid-distance between the pit and lower part of the asymmetrica-Norigondolella sp. Subzone of the posterior margin; secondary keels extend close to the postero-lateral primitia Zone. Mazza et al. (2012a) record their elements from Lower corners. Lacian? strata, but show they are entirely restricted to the “boundary Comparisons: Primatella vanlierae is similar to the present species interval” (op. cit., fig. 2). but the posterior margin of those P1 elements is straight or only slightly Primatella circulare sp. nov. indented. Other posteriorly expanded species such as P. stanleyi and Figure 63. 4-22 P. triangulare lack the posterior sinus. Several other species are characterized by a bifid posterior margin, but they lack anterior nodes 2014 Primatella sp. nov. 4 - Balini et al., fig. 11.m1-3. or have only poorly developed ones (see below). Derivation of name: Refers to the circular outline of the posterior Remarks: This is a rather variable form in terms of platform outline, platform. and the anterior node and carina formation. The carina may terminate Holotype: GSC 132964, Figure 63. 17-19. well in front of the posterior margin, or may have a node developed close to that margin. The anterior nodes may be sharp or blunt, discrete Type stratum: Bed 18h, within the Pardonet Formation. or partly coalescing. However, the posterior sinus is a distinctive Age: Middle subdivision of the parvus Subzone of the primitia Zone. unifying feature. 91

Figure 63. 1-3. Quadralella? sp. nov. A. GSC 132957, sample 17. 4-22. Primatella circulare sp. nov. 4, 5. GSC 132959, sample 21d. 6, 7. GSC 132960, sample 20. 8-10. GSC 132961, sample 21d. 11-13. GSC 132962, sample 07-22. 14-16. GSC 132963, sample 18f. 17-19. GSC 132964, sample 18h. Holotype. 20-22. GSC 132965, sample PHE-23. Pardonet Hill east. Scale bar = 200 microns.

Diagnosis: The platform of the P1 element is divided into two subequal Type stratum: Bed 21d, within the Pardonet Formation. parts by a strong constriction: an anterior half with straight margins Age: Upper subdivision of the parvus Subzone of the primitia Zone. bearing 2-4 discrete and sharp denticles, and a posterior half with a subcircular outline. The blade is between ½-⅓ element length and Diagnosis: An elongate Primatella with a posterior carina composed descends onto the platform as a row of four nodes, the last of which of two well separated carinal nodes, the posterior of which lies well is the largest and occurs a little posterior of the constriction; a small in front of the posterior end of the platform, with the consequent accessory node may occur on an otherwise unornamented posterior development of a broad platform brim (Orchard, 2013). platform brim. A pit occurs beneath the constricted part of the platform, Comparisons: Primatella conservativa is closely allied to P. primitia a little posterior of the midpoint, and slightly displaced anteriorly (q.v.) but the posterior nodes of the carina differ; P. elongata differs within the keel. in its longer carina. Each of these species has longer and narrower Comparisons: The configuration of the carina and cusp of the P1 platforms than P. mersinensis. element of Primatella circulare most closely resembles those of P. Remarks: Primatella conservativa was included in Epigondolella orchardi, but the posterior platform of the latter is subrectangular primitia by Mosher (1970, 1973) but the latter species, in the restricted rather than rounded. Other Primatella species with a medial platform sense of Kozur and Moix (in Moix et al., 2007), is far less common at constriction are longer and have differing carinas and posterior platform Black Bear Ridge, contrary to the assumption of Mazza et al. (2012a). outlines. Stratigraphic occurrence: Abundant at Black Bear Ridge, the species Stratigraphic occurrence: Common in the parvus Subzone through is the first of the “typical”Primatella species to appear near the base of lower asymmetrica-Norigondolella sp. Subzone of the primitia Zone; the primitia Zone, and it ranges through the remainder of the zone. The rare higher in the latter subzone. two specimens illustrated by Mosher (1970) and included here came Primatella conservativa Orchard from matrix of the Carnian Macrolobatus Zone of Pardonet Hill (GSC Figure 64. 1-23. cur. no. 64628), whereas those illustrated by Orchard (1983) came from the Norian Kerri Zone. 1970 [p] Epigondolella primitia n. sp. - Mosher, p. 740-41, pl. 110, figs. 10, 13, 16, 17 (only). Primatella elongata sp. nov. 1973 [p] Epigondolella primitia Mosher - Mosher, p. 161, pl. 18, figs. Figure 65. 1-21. 7, 8, 9, 10 (only). Derivation of name: Refers to the elongate posterior carina. 1983 Epigondolella primitia Mosher - Orchard, p. 178-9, fig. 15A- C. Holotype: GSC 132983, Figure 65. 16-18. 2013 Primatella conservativa sp. nov. - Orchard, p. 455, fig. 4.1- Type stratum: Bed 07-13a, within the Pardonet Formation. 4.3. 2013 Primatella conservativa Orchard - Carter and Orchard, fig. 7. Age: The angusta-dylani Subzone of the primitia Zone. 1-3. Diagnosis: A species of Primatella characterized by P1 elements with 2014 Primatella conservativa Orchard - Balini et al., fig. 10 a1-3, an elongate platform that abruptly narrows in its posterior half. The b1-3. anterior platform bears 2-4, rarely up to 6 discrete, sharp nodes on Holotype: GSC 132610, Figure 64. 21-23. each margin. The posterior platform is unornamented at its margins, 92 93

Figure 65. 1-21. Primatella elongata sp. nov. 1-3. GSC 132978, sample 21f. 4-6. GSC 132979, sample 18h. 7-9. GSC 132980, sample 07-18. 10-12. GSC 132981, sample 21d. 13-15. GSC 132982, sample 18. 16-18. GSC 132983, sample 07-13a. Holotype. 19-21. GSC 132984, sample 17b. Scale bar = 200 microns. which are subparallel laterally and squared off posteriorly. The blade is Figure 66. 1-18, ?19-21 between ½ and ⅓ element length and passes onto the platform as a row of carinal nodes (commonly 7) that extend close to the posterior end Derivation of name: Named for Frank McLearn, pioneering Triassic of the platform. In larger specimens, the posteriormost node is smaller biostratigrapher in northeastern British Columbia. than those to the anterior, one of which (commonly the penultimate Holotype: GSC 132987, Figure 66. 7-9. one) is distinctly larger than all others. A very narrow posterior platform brim is present. The lower surface has a broad keel and a subcentral pit Type stratum: Bed 13c, within the Pardonet Formation. far anterior to the posterior end of the keel, which may have a straight, Age: The angusta-dylani Subzone of the primitia Zone. oblique, or bifid termination. Diagnosis: An elongate species of Primatella characterized by a P1 Comparisons: These elements were formerly included in a broad platform with a medial constriction separating a marginally ornate concept of the species primitia, but it is distinguished from other similar anterior platform from an initially narrower and, with growth, a elements by the longer posterior carina. Unlike Primatella primitia and posteriorly expanded posterior platform. The nodes or denticles on P. conservativa, P. elongata possesses a carina that extends close to the the anterior platform margins are generally well differentiated but are posterior edge of the platform. In this respect, it resembles Acuminatella apically rounded and may be partly fused and irregularly developed. angusta but that species has a less developed anterior platform with The posterior platform is subrectangular in early growth stages but fewer, often smaller nodes, a narrower posterior platform, and a high usually expands towards the posterior margin with growth. The free and fused posterior carina that commonly extends to the narrower, blade is approximately ⅓ total element length and passes onto the pointed end of the posterior platform. platform as 5-7 carinal nodes that may terminate in a large cusp in Stratigraphic occurrence: Very common in the angusta-dylani front of a wide platform brim, or as 1-3 partly fused and raised nodes Subzone through the asymmetrica-Norigondolella sp. Subzone of the that extend close to the posterior margin. The pit is located posterior of primitia Zone. the platform midlength and is generally only slightly shifted anteriorly Primatella mclearni sp. nov. within the keel. Figure 64 (facing page). 1-23. Primatella conservativa Orchard. 1-3. Comparisons: This, the oldest species assigned to Primatella in the GSC 132966, sample 28. 4-6. GSC 132967, sample 17a. 7-9. GSC present collections, differs from all younger species with a similar 132968, sample 21h. 10, 11. GSC 132969, sample 15. 12-14. GSC medially constricted platform in its relatively poorly developed anterior 132970, sample 20f. 15-17. GSC 132971, sample 18h. 18-20. GSC nodes and the relatively posterior pit position. In younger species, 132972, sample 21d. 21-23. GSC 132610, sample 21d. Holotype. 24- anterior nodes are invariably higher, more discrete, and apically sharp, 44. Primatella primitia (Mosher). 24-26. GSC 132973, sample 18f. 27- and the pit is more anterior relative to both the platform and keel 29. GSC 132974, sample 13a. 30-32. GSC 132975, sample 18e. 33-35. terminations. Parapetella lanei has a similar platform shape but lacks GSC 132976, sample 10/04. 36-38. GSC 25051. Holotype. GSC cur. anterior ornament. no. O-64654, Brown Hill. 39-41. GSC 132614, sample 20b (not 18e, Remarks: Elements combined here show variation in posterior carina as in Orchard 2013). 42-44. GSC 132977, sample 20d. Scale bar = 200 development, including elements with long carinas extending close to microns. 94

Figure 66. 1-9, ?10-12, 13-18, ?19-21. Primatella mclearni sp. nov. 1-3. GSC 132985, sample 5. 4-6. GSC 132986, sample 04-25. 7-9. GSC 132987, sample 13c. Holotype. ?10-12. GSC 132988, sample 4. 13-15. GSC 132989, sample 14. 16-18. GSC 132990, sample 17a. ?19-21. GSC 132991, sample 18f. Advanced form with more anterior pit. Scale bar = 200 microns. the posterior margin and others with wide posterior platform brims. 2014 Primatella mersinensis (Kozur and Moix) - Balini et al., fig. Younger species of Primatella are distinguished on the basis of this 10 e1-3. feature, and variation in P. mclearni may reflect early diversification of the root stock from which a variety of younger species developed. Description: The P1 element has a subrectangular platform with several Questionably included here are the stratigraphically lowest short and discrete nodes or denticles on the anterior lateral margins, specimen (Fig. 66. 10-12), which has an atypical posterior carina, and unornamented posterior platform margins. Generally, a relatively and the stratigraphically youngest specimen (Fig. 66. 19-21), which small subcentral cusp is separated from 2, rarely 3 larger discrete nodes is exceptional in having low rounded anterior nodes and a keel that that occur to its posterior. The pit lies below the center of the platform, extends far posterior of the pit. The latter may represent an advanced anteriorly displaced within a keel that is posteriorly bifurcated. morphotype, or reflect a gerontic morphology. Comparisons: The P1 elements of Primatella conservativa, P. elongata, and P. primitia all differ from the present species in having relatively Stratigraphic occurrence: Questionable in the topmost spenceri longer and narrower platforms, whereas P. stanleyi and P. orchardi Subzone of the samueli Zone, but abundant in the sagittalis-beattyi differ in having a platform constriction. The elements are closest in Subzone through the acuminata-prominens Subzone, and questionable shape to P. subquadrata, but P. mersinensis differs in having additional in the lower subdivision of the parvus Subzone of the primitia Zone. posterior carinal nodes. Late growth stages of P. mersinensis maintain Primatella mersinensis (Kozur and Moix) a rectangular shape, in contrast to P. triangulare in which the posterior Figure 67. 19-24; 25-36, Figure 68. 1-12 platform is expanded into a triangular shape. A Norian example of 1989 Epigondolella primitia Mosher - Orchard, in Carter et al., pl. the present species (Fig. 67. 31-33) has acutely angled postero-lateral 1, fig. 16. corners, a feature more commonly seen in the alpha morphotype of 2000 Metapolygnathus primitius (Mosher) - Carter and Orchard, Epigondolella quadrata, which, however, has much larger anterior pl. 1, fig. 8. denticles. 2007 Metapolygnathus mersinensis n. sp. - Kozur and Moix, in Remarks: The holotype of Primatella mersinensis, perhaps atypically, Moix et al., p. 293, pl. 1, figs. 14a-c, 15a-b. 2010 “Metapolygnathus communisti B” Krystyn - Mazza et al., pl. Figure 67 (facing page). 1-18. Primatella subquadrata sp. nov. 3, figs. 4a-c. 1-3. GSC 131260, sample 10/03a. 4-6. GSC 131261, sample 07-22. 2012 [p] Metapolygnathus mersinensis Kozur and Moix - Mazza et 7-9. GSC 131262, sample PHE-23. Pardonet Hill east. 10-12. GSC al.(b) , p. 116-17, pl. 4, figs. 6a-c (only). [fig. 4= P. aff. 131263, sample 20c. 13-15. GSC 131264, sample 21g. 16-18. GSC asymmetrica; figs. 7a-c, 9a-c = P. subquadrata; 5, 8, ?10 = 131265, sample 18a. Holotype. 19-24. Primatella mersinensis (Kozur Quadralella spp.]. and Moix), alpha morphotype. 19-21. GSC 131266, GSC cur. 2012 Metapolygnathus cf. primitius (Mosher) - Mazza et al., (b), p. no. O-64654, Brown Hill. 22-24. GSC 131267, sample 17c. 25-36. 117, pl. 8, fig. 12 (=M. mazzai sensu Karadi et al., 2013). Primatella mersinensis (Kozur and Moix), beta morphotype. 25-27. 2013 Primatella mersinensis (Kozur and Moix) - Carter and GSC 131268, sample 21f. 28-30. GSC 132612. GSC cur. no. O-64654, Orchard, fig. 7. 7-9. Brown Hill. 31-33. GSC 131269, sample 21h. 34-36. GSC 131270, sample 21g. Scale bar = 200 microns. 95 96

Figure 68. 1-12. Primatella mersinensis (Kozur and Moix), beta morphotype. 1-3. GSC 131271, sample 31. 4-6. GSC 131272, sample 21f. 7-9. GSC 131273, sample 17c. 10-12. GSC 131274, sample 07-22. 13-21. Primatella triangulare sp. nov. 13-15. GSC 131275, sample 21g. 16-18. GSC 131276, sample PHE-24. Pardonet Hill east. 19-21. GSC 131277, sample PHE-24. Holotype. Pardonet Hill east. Scale bar = 200 microns. 97 has a continuation of the carina as a low ridge extending to one postero- in the sagittale-beattyi Subzone through the acuminata-prominens lateral margin. This appears to be yet another variation in posterior carina Subzone of the primitia Zone. The beta morphotype is uncommon configuration, although comparable forms have not been observed in in the acuminata-prominens Subzone through the asymmetrica- the present material. Kozur and Moix (in Moix et al., 2007) illustrated Norigondolella sp. Subzone of the primitia Zone. only two specimens of this species from Turkey, neither of them in full upper view, so it is difficult to appreciate the character and variability Primatella orchardi (Kozur) of the species. Neither figured specimen has large anterior nodes, but Figure 70. 1-3?, 4-26 rather rounded low nodes in one case and short, sharper denticles in the 2003 [p] Epigondolella orchardi n. sp. - Kozur, p. 69-70, pl. 1, figs. second; the breadth of the platform is equal throughout its length and 1a-c, 7a-b (only). it does not have a constriction. The present material includes similar 2013 Primatella orchardi (Kozur) - Carter and Orchard, fig. 7. 4-6. elements that have more pronounced anterior nodes, and rare large P1 2014 Primatella orchardi (Kozur) - Balini et al., fig. 10 h1-3 (notP. specimens with elaboration of the posterior margin. sp. nov. 7, =fig. 10 g1-3). Karadi et al. (2013) introduced Metapolygnathus mazzai based on material from Csövar, Hungary and selected a specimen from Pizzo Diagnosis: The P1 platform elements have a submedial constriction, Mondello assigned to M. cf. primitius as the holotype (Mazza et al., particularly in early growth stages, that separates 2-4 well developed, 2012b, pl. 8, fig. 12). This specimen corresponds in most respects to round to laterally enlarged, pointed to round-topped nodes on each the holotype of M. mersinensis, which Karadi et al. (2013) regarded as anterior platform margin from an unornamented posterior platform. the ancestor of their species. Neither illustrated specimen of M. mazzai The posterior platform is typically subrectangular with a breadth equal from Hungary is clearly the same as the holotype. At the moment, to or greater than that of the anterior platform. The carina terminates in there is no clear distinction between M. mazzai and M. mersinensis as a large and discrete terminal cusp opposite or slightly posterior to the represented by their holotypes. platform constriction, and in front of a broad platform brim. The pit lies Two morphotypes of Primatella mersinensis are distinguished beneath or slightly posterior of the centre of the platform, opposite the in this study on the basis of the configuration of the posterior carina: constriction, and shifted anteriorly with respect to the keel, which may these have direct counterparts in P. stanleyi. The uncommon alpha be bifurcated in larger specimens. morphotype (Fig. 67. 19-24) has a pair of partly fused posterior carinal Comparisons: Primatella stanleyi has a similar platform but has nodes, and the more common beta morphotype (Fig. 67. 25-36, Fig. additional posterior carinal denticles, whereas P. subquadrata has a 68. 1-12) has 2-3 separated nodes. similar carina but lacks a distinct platform constriction. Elements of the Mazza et al. (2012b) described and illustrated a wide variety of older Carnepigondolella pseudodiebeli beta morphotype are similar, elements under this species, including two morphotypes in addition but they have a less constricted, relatively short platform, a more to a central one. They differ in relative platform shape, carina length, posteriorly situated pit, and a shorter blade. and anterior node formation. Most of these specimens are excluded Remarks: In the original diagnosis for this species, it was noted from the current species because they lack either clearly differentiated that lateral margins of the platform were parallel or subparallel, but and pointed anterior nodes, or a posterior carina (see Primatella also an “indistinct constriction at the beginning of the posterior third subquadrata). of the platform is often present.” Kozur (2003) illustrated only three Stratigraphic occurrence: The beta morphotype is abundant, and the specimens, each with a different platform outline: the P1 specimen from alpha morphotype is rare, in the angusta-dylani Subzone through the Turkey (op. cit., fig. 7) has a distinct constriction about its platform asymmetrica-Norigondolella sp. Subzone of the primitia Zone. The midlength, whereas those from Slovakia have an indistinct constriction holotype was described from the orchardi Zone in the Kocatabur Block (op. cit., fig. 1, the holotype) or evidently lack one (op. cit., fig. 2). The of the Mersin Melange in southeast Turkey (Moix et al., 2007), where holotype also has a relatively longer platform than the other specimens it is associated with an uppermost Carnian association (Mazza et al., illustrated by Kozur. In the British Columbian material, there are also 2012b). The various elements reported by Mazza et al. (2012b) and examples of each of the European forms and they apparently have combined here in Primatella mersinensis have various ranges within slightly different stratigraphic occurrences. For this reason, elements the total range of upper Tuvalian to lower Lacian. with a distinctive constriction are separated from those that have Primatella oblonga sp. nov. relatively shorter platforms of constant width, which are assigned to Figure 69. 1-6; 7-12 Primatella subquadrata. Specimens with shorter and posteriorly expanded platforms, and 2014 Primatella sp. nov. 8 - Balini et al., fig. 11.d1-3. shorter blades assigned by Mazza et al. (2012b, pl. 2, figs. 1a-c. 2a- c) to P. orchardi are excluded from this species: their stratigraphic Derivation of name: Refers to the oblong outline of the P1 platform position suggests they may be representative of Carnepigondolella element. pseudodiebeli beta morphotype (q.v.). One specimen (Fig. 70. 1-3), Holotype: GSC 131281, Figure 69. 10-12. questionable included here, differs in having slightly ornate posterior platform margins and an inconspicuous terminal cusp. Note that the Type stratum: Bed 17b, within the Pardonet Formation. illustration of Primatella orchard in Balini et al. (2014, fig. 10 h1-3) Age: The acuminata-prominens Subzone of the primitia Zone. was swapped with that of Primatella sp. nov. 7 (=P. triangulare, op. cit., Diagnosis: The P1 elements of this species are characterized by an fig. 10 g1-3). elongate, oblong-shaped platform with a rounded posterior outline. The Stratigraphic occurrence: Abundant in the angusta-dylani Subzone anterior half of the platform bears 4-5 well developed marginal nodes through the asymmetrica-Norigondolella sp. Subzone of the primitia that tend to be sharply terminated and higher to the anterior and lower Zone. and more rounded to the posterior. Three widely spaced carinal nodes of variable size are commonly developed posterior to the cusp, in front Primatella ovale sp. nov. of a narrow posterior platform brim. The pit is located at or a little Figure 71. 21-23, ?24-26 posterior to the platform midpoint, and anterior to the narrowly bifid or Derivation of name: Refers to the oval shaped platform of the P1 rounded posterior keel margin. element. Comparisons: These elements have a similar carinal configuration Holotype: GSC 131308, Figure 71. 21-23. to Primatella elongata but they differ in having a rounded posterior margin. The similar elements of P. rotunda have a shorter, broader Type stratum: Bed 18h, within the Pardonet Formation. platform, and a shorter carina. Age: Lower subdivision of the parvus Subzone of the primitia Zone. Remarks: The oldest examples of this species have well differentiated Diagnosis: The P1 element has an oval shaped platform that is broadest but low rounded nodes on the anterior margins, whereas younger in the anterior, slowly tapered to the posterior, and terminated in a examples have sharp anterior denticles that extend progressively farther narrowly to broadly rounded posterior margin. The anterior platform to the posterior. These forms are differentiated, respectively, as alpha margins bear 2-4 discrete and sharp denticles, whereas the posterior morphotype (Fig. 69. 1-6) and beta morphotype (Fig. 69. 7-12). is unornamented. The free blade is about ⅓ total element length and descends onto the platform as 3-4 separated carinal nodes, the last one Stratigraphic occurrence: The alpha morphotype is uncommon of which is conspicuous and well in front of the posterior margin. The 98

Figure 69. 1-6. Primatella oblonga sp. nov., alpha morphotype. 1-3. GSC 131278, sample 07-7. 4-6. GSC 131279, sample 14a. 7-12. Primatella oblonga sp. nov., beta morphotype. 7-9. GSC 131280, sample 17b. 10-12. GSC 131281, sample 17b. Holotype. 13-21. Primatella rotunda sp. nov., alpha morphotype. 13-15. GSC 131283, sample 17. 16-18. GSC 131284, sample 17. Holotype. 19-21. GSC 131285, sample 07-18. 22-30. Primatella rotunda sp. nov., beta morphotype. 22-24. GSC 131286, sample 21h. 25-27. GSC 131287, sample 21g. 28-30. GSC 131288, sample 07-18. Scale bar = 200 microns.

Figure 70 (facing page). 1-3?, 4-26. Primatella orchardi (Kozur). 1-3. GSC 131289, sample 15. 4-6. GSC 131290, sample 21h. 7-9. GSC 131291, sample 07-22. 10-12. GSC 131292, sample 17a. 13-15. GSC 131293, sample 18f. 16-18. GSC 131294, sample 22. 19-21. GSC 132611, sample 21h (sample 3 in Orchard, 2013 is incorrect). 22-24. GSC 131295, sample 10/03. 25, 26. GSC 131296, sample 18e. 27-32. Primatella stanleyi sp. nov., alpha morphotype. 27-29. GSC 131297, sample 26. 30-32. GSC 131298, sample 18. 33-44. Primatella stanleyi sp. nov., beta morphotype. 33-35. GSC 131299, sample 16. 36-38. GSC 131300, sample 21f. 39-41. GSC 131301, sample 18f. 42-44. GSC 132591, sample PHE-23. Holotype. Pardonet Hill east. Scale bar = 200 microns. 99 100 101 Figure 71 (facing page). 1-14. Quadralella stephanae (Orchard). 1, Primatella posteroglobosa sp. nov. 2. GSC 131302, sample 07-8. 3-5. GSC 131303, sample 11c. 6-8. GSC Figure 73. 1-19 131304, sample 8. 9-11. GSC 131305, sample 12. 12-14. GSC 132603, Derivation of name: Refers to the expanded globular-like outline of sample 5. 15-20. Primatella aff. ovale. 15-17. GSC 131306, sample the posterior platform of the P1 element. 07-12c. 18-20. GSC 131307, sample 13a. 21-23, ?24-26. Primatella ovale sp. nov. 21-23. GSC 131308, sample 18h. Holotype. ?24-26. Holotype: GSC 131331, Figure 73. 14-16. GSC 131309, sample 21f. 27-47. Primatella ex gr. pseudoechinata Type stratum: Bed 17b, within the Pardonet Formation. (Kozur). 27-29. GSC 131310, sample 07-22. 30-32. GSC 131311, sample PHE-25. Pardonet Hill east. 33-35. GSC 132613, sample 17. Age: The acuminata-prominens Subzone of the primitia Zone. 36-38. GSC 131312, sample PHE-23. Pardonet Hill east. 39-41. GSC Diagnosis: The P1 element has a slender platform with a strong medial 131313, sample 10/04. 42-44. GSC 131314, sample 10/04. 45-47. GSC constriction, anterior of which there are 2-3 well developed, apically 131315, sample 10/04. 48-53. Primatella rhomboidale sp. nov. 48-50. sharp marginal nodes on each lateral margin. Posteriorly, the platform GSC 131316, sample 21f. Holotype. 51-53. GSC 131317, sample 18d. is narrower than the anterior platform and has a lachrymiform outline. Scale bar = 200 microns. The free blade is about ⅓ element length and descends onto the pit occupies a submedial position under the platform within a keel that platform as 2-3 small anterior carinal nodes and then commonly as 3 extends to its posterior. larger, partly fused posterior nodes that almost extend to the posterior platform margin, from which they are separated by a narrow brim. The Comparisons: These P1 elements resemble those of Primatella ex gr. pit lies beneath the platform midlength, well in front of the end of the pseudoechinata (q.v.), but the former have a longer, narrower, and more keel. differentiated posterior platform. Comparisons: The older species and probable precursor Primatella Remarks: A large specimen (Fig. 71. 24-26) questionably included medioconstricta has a similar though less bulbous posterior platform, here has a posterior platform and keel that is longer than is typical. less well developed anterior nodes, and a more posteriorly located pit. The anterior denticulation is atypically irregular, but the conspicuous Similar sized elements of P. conservativa have a shorter, subrectangular terminal node is like that of the holotype. posterior platform, whereas P1 elements of P. vanlierae are much more Stratigraphic occurrence: Uncommon in the parvus Subzone and laterally expanded at their posterior end. Acuminatella species have a asymmetrica-Norigondolella sp. Subzone of the primitia Zone. much narrower, often pointed and unexpanded posterior platform, and a strong posterior carina. Primatella aff. ovale Figure 71. 15-20 Stratigraphic occurrence: Very common in the angusta-dylani Subzone and into the asymmetrica-Norigondolella sp. Subzone of the Remarks: These elements differ from Primatella ovale in having primitia Zone. apically rounded anterior nodes, and a pit that occupies a more posterior position in the keel. Primatella primitia (Mosher) Figure 64. 24-44 Stratigraphic occurrence: Rare in the angusta-dylani Subzone of the primitia Zone. 1970 [p] Epigondolella primitia n. sp. - Mosher, p. 740-41, pl. 110, figs. 8, 1,1 12 (only). Primatella permica (Hayashi) 1973 Epigondolella primitia Mosher - Mosher, p. 161, pl. 18, figs. Figure 72. 1-22 1, 2, 11 (only). 1968 Gladigondolella abneptis var. permica var. nov. - Hayashi 1991 Metapolygnathus primitia Mosher - Orchard (b), p. 317-8, (a), p.69, pl. 2, figs. 9a-c. pl. 1, figs. 13-15. 1968 Gladigondolella abneptis permica n. subsp. - Hayashi (b), 2013 Primatella primitia (Mosher) - Orchard, p. 453-4, fig. 4. 13- p.11. 15. 2003 Epigondolella permica (Hayashi) - Channel et al., fig. A1. 46, 2013 Primatella primitia (Mosher) - Carter and Orchard, fig. 7. 10- 54, 54, 56. 12. 2014 Primatella sp. nov. 1 - Balini et al., fig. 10 i1-3. Holotype: GSC 25051, re-illustrated here in Figure 64. 36-38. Diagnosis: The P1 elements of this Primatella species are characterized Type stratum: Upper Kerri Zone within Pardonet Formation. by irregularly developed marginal nodes on both the lateral and posterior platform margins, although these are never as strongly developed as Type locality: GSC cur. no. O-64654, Brown Hill. those on the anterior margins. The essentially subrectangular platform Description: The P1 element of this species is relatively long, with 2-5 has an irregular shape due to the development of low, generally anterior denticles on each margin, a medial constriction (more evident in irregular marginal nodes that project outwards from the lateral and earlier growth stages), an unornamented rectangular posterior platform, posterior margins. The blade is between ½ and ⅓ total element length and carinal nodes that extend onto the posterior platform. Three closely and passes into 4 discrete carinal nodes that are separated from the spaced carinal nodes lie posterior of a gap that separates them from the posterior platform margin by a brim. The pit is submedial in position, cusp; a tiny fourth accessory node may occur to the posterior. A broad and progressively anteriorly shifted in the keel. platform brim separates the posterior carina from the platform margin. Comparisons: The P1 element of this species differs from that of The posterior platform broadens with growth, and any constriction P. rhomboidale in its irregular posterior ornament and generally is overgrown so as to produce elements of uniform breadth. The pit asymmetric shape. The overall shape may resemble rectangular species generally occupies a position beneath the platform midlength or a little like Primatella mersinensis, but the posterior marginal ornamentation to the anterior in mature specimens, and is anteriorly shifted within the sets them apart. Similarly, this difference also distinguishes smaller keel, which may have a straight, oblique, or bifid termination. elements, like the holotype, that have a medial constriction like that Comparisons: Primatella primitia differs from all other species of seen in P. orchardi and P. stanleyi. the genus in having three closely spaced carinal nodes posterior of the There are many younger species of Epigondolella with posterior cusp. The alpha morphotypes of both P. stanleyi and P. mersinensis nodes, but they all have larger and higher anterior nodes, as well as have two closely adjacent carinal nodes, but the former has a persistent an upturned posterior platform and less compact microreticulation constriction, and the latter has a shorter subrectangular platform. on the platform. This species, which appears to be the first taxon to Primatella conservativa and P. elongata have similar platform develop posterior ornament since the extinction of Carnepigondolella, proportions but posterior carinal nodes are discrete and, in the case of is followed successively by similar but more ornate Epigondolella aff. P. elongata, the posterior carina is more developed and extends close to uniformis and E. uniformis in the Norian. the posterior edge of the platform. Stratigraphic occurrence: Very common in the acuminata-prominens Remarks: The present concept of this species is very restricted Subzone through the asymmetrica-Norigondolella sp. Subzone of compared with that of Mosher (1970, 1973), whose holotype is the only the primitia Zone. The Japanese holotype came from a mixed fauna specimen he illustrated that has the characteristic posterior carina; its extracted from chert. Slovakian examples come from the late Tuvalian pit is also located more to the anterior than is typical. Such specimens (Channel et al., 2003). 102

Figure 72. 1-22. Primatella permica (Hayashi). 1-3. GSC 131318, sample 18. 4-6. GSC 131319, sample 21a. 7, 8. GSC 131320, sample 20. 9, 10. GSC 131321, sample 21. 11-13. GSC 131322, sample 21d. 14-16. GSC 131323, sample 17c. 17-19. GSC 131324, sample 22. 20-22. GSC 131325, sample 20. Scale bar = 200 microns. are rare in the large collections from British Columbia where elements nov. B and P. sp. nov. C differ in their anterior parapets and posterior now referred to Primatella conservativa are far more common. The two constriction. The diminutive P1 element of Metapolygnathus echinatus may be indistinguishable in early growth stage. (Hayashi) (=Carnepigondolella echinatus sensu Carter and Orchard, Stratigraphic occurrence: Uncommon in the angusta-dylani Subzone 2013) has a much longer blade, smaller platform, and an anterior pit. through the asymmetrica-Norigondolella sp. Subzone of the primitia Remarks: Since Primatella pseudoechinata was first described from Zone. Mosher’s (1970) illustrated specimens of “Epigondolella” the Late Carnian Macrolobatus Zone of northern Hungary (Kovacs and primitia both from the Late Carnian and Early Norian but the holotype Kozur, 1980), it has been broadly interpreted and often misidentified. and only typical specimen came from Subzone II of the Kerri Zone. The For example, Orchard (2007a, b) incorrectly assigned specimens to this species is also known from Haida Gwaii (Carter and Orchard, 2013). species that are now referred to new species of Carnepigondolella. Primatella ex gr. pseudoechinata (Kozur) The uncommon representatives of the pseudoechinata group Figure 71. 27-47 identified here differ in detail and further subdivision may be warranted. A specimen from Pardonet Hill east (e.g., Fig. 71. 36-38) is closest to 1989 Epigondolella pseudoechinata n. sp. - Kozur (b), p. 430, 431. the symmetrical holotype. Other elements have more variable posterior 1991 Metapolygnathus pseudoechinatus (Kozur) - Beyers and platforms and anterior nodes, and the youngest examples include Orchard, pl. 4, fig. 20. specimens (Fig. 71. 45-47) with a distinct parapet rather than denticles 2000 Metapolygnathus pseudoechinatus (Kozur) - Carter and on one anterior margin. Orchard, pl. 1, fig. 3. 2013 Primatella pseudoechinata (Kozur) - Carter and Orchard, fig. Stratigraphic occurrence: Questionably in the angusta-dylani 7. 19-21. Subzone, and uncommon in the acuminata-prominens Subzone through the asymmetrica-Norigondolella sp. Subzone of the primitia Zone. The Description: The P1 elements combined here have in common a element closest to the holotype is from the asymmetrica-Norigondolella relatively short, equidimentional platform that is broadest in the anterior sp. Subzone of Pardonet Hill. The element illustrated from Haida Gwaii part and tapers to a broadly rounded or truncated posterior margin. also occurs in the Lower Norian (Carter and Orchard, 2000). Each anterior platform margin bears 2-3 discrete nodes or denticles, whereas the posterior part is unornamented. The free blade is between Primatella rectangulare sp. nov. ⅓ and ½ of the total element length and descends on to the platform as Figure 74. 1-12 2-4 discrete carinal nodes, the last of which is in front of the posterior Figure 73 (facing page). 1-19. Primatella posteroglobosa sp. nov. platform margin. The pit occupies a position beneath or posterior to the 1-3. GSC 131326, sample 18e. 4-6. GSC 131327, sample 17. 7, 8. platform midlength, and the keel extends only a short distance beyond GSC 131328, sample 16. 9, 10. GSC 131329, sample 13a. 11-13. GSC it where it may be rounded, straight, pointed, or bifurcate. 131330, sample 14a. 14-16. GSC 131331, sample 17b. Holotype. 17-19. Comparisons: The squat, equidimentional platforms of these elements GSC 131332, sample 17. 20-43. Primatella vanlierae sp. nov. 20-22. distinguish them from most other species of Primatella. They are GSC 131333, sample 24. 23-25. GSC 131334, sample 29. 26-28. GSC shorter and posteriorly broader than P. ovale. Similarly shaped platform 131335, sample 18h. Holotype. 29-31. GSC 131336, sample 07-12a. elements of Quadralella stephanae lack discrete anterior platform 32-34. GSC 131337, sample 18f. 35-37. GSC 131338, sample 17b. denticles, and those of Carnepigondolella spenceri have a posteriorly 38-40. GSC 131339, sample PHE-23. Pardonet Hill east. 41-43. GSC broader platform and smaller anterior nodes. Both Parapetella sp. 131340, sample PHE-23. Pardonet Hill east. Scale bar = 200 microns. 103 104

Figure 74. 1-12. Primatella rectangulare sp. nov. 1-3. GSC 131341, sample 5. 4-6. GSC 131342, sample 07-12. 7-9. GSC 131343, sample 8. Holotype. 10-12. GSC 131344, sample 04-11B. Scale bar = 200 microns. Derivation of name: Refers to the rectangular outline of the P1 element. Primatella rotunda sp. nov. Holotype: GSC 131343, Figure 74. 7-9. Figure 69. 13-21; 22-30 Type stratum: Bed 8, within the Pardonet Formation. 1982 [aff.] Neogondolella nodosa (Hayashi) - Koike, p. 24, pl. 2, figs. 7-9 (only). Age: The sagittale-beattyi Subzone of the primitia Zone. 2010 Carnepigondolella nodosa B - Mazza et al., pl. 1, fig. 5a-c. Diagnosis: A species of Primatella in which the P1 element is 2012 [?p] Carnepigondolella tuvalica n. sp. - Mazza et al.(b), p. 100- subrectangular in outline and bears poorly differentiated and irregular 03, pl. 3, figs. 6a-c, 7a-c (only). nodes along the anterior platform margins, and a generally inornate 2014 Primatella sp. nov. 2 - Balini et al., fig. 11.i1-3, j1-3. posterior platform. The blade is ~⅓ element length and continues as Derivation of name: Refers to the broad, rounded outline of the P1 a row of carinal nodes that terminates in a larger cusp or node-pair platform element. located between the platform center and its posterior margin; there is a broad platform brim. The pit is located posterior of midlength, and is Holotype: GSC 131284, Figure 69. 16-18. slightly displaced to the anterior within the keel. Type stratum: Bed 17, within the Pardonet Formation. Comparisons: The anterior marginal nodes of the P1 element of Age: The acuminata-prominens Subzone of the primitia Zone. this species are similar to those of the contemporaneous Primatella mclearni, but the present species is relatively shorter and broader, and Diagnosis: The P1 elements have broad platforms with biconvex to lacks a well-defined medial platform constriction. In common with parallel lateral margins and well-rounded postero-lateral margins. the former species, P. rectangulare lacks the regular and pronounced The anterior margins bear variably and unevenly developed, apically anterior denticles of younger Primatella species, and has a more rounded to pointed nodes that may be discrete or coalescing. The free posteriorly located pit. blade is about ⅓ unit length, but may be relatively longer in small specimens and shorter in some large specimens in which the platform Stratigraphic occurrence: Very common in the sagittale-beattyi flanges extend far to the anterior. The carinal denticles are generally Subzone through angusta-dylani Subzone of the primitia Zone. discrete and extend on to the posterior platform where a terminal or Primatella rhomboidale sp. nov. subterminal cusp of variable size lies well in front of a wide posterior Figure 71. 48-53 platform brim. The pit varies between a medial and posterior position within the furrowed keel. Derivation of name: Refers to the rhomb-shaped platform P1 element. Comparisons: These elements have a posteriorly rounded shape that Holotype: GSC 131316, Figure 71. 48-50. readily distinguishes them from the more commonly subrectangular and Type stratum: Bed 21f, within the Pardonet Formation. constricted species of Primatella. Within the genus, a rounded posterior margin characterizes the P1 elements of P. circulare, which has a Age: Upper subdivision of parvus Subzone of the primitia Zone. strongly constricted platform, and of P. oblonga, which is relatively Diagnosis: The P1 element has a rhomb-shaped platform, 3-4 well longer, narrower, and has a longer carina. The older Carnepigondolella developed platform denticles on each anterior margin, 1-2 more milanae has a similar platform shape, but it has smaller, more discrete rounded nodes on each posterior lateral margin, and a low node forming anterior denticles. Some Kraussodontus and Parapetella species have a pointed posterior margin that is aligned with the carina to the anterior. rounded outlines, but they lack the anterior nodes. Metapolygnathus The blade is ~⅓ element length and passes into a row of 4-5 carinal multinodosus Noyan has a more rectangular platform and combines nodes, the last of which is a more separated cusp. The pit lies posterior “primitive” anterior nodes with an “advanced” anterior pit. of midlength and near the posterior end of a bifid keel. Remarks: This species is assigned to Primatella based on its Comparisons: The posterior marginal ornament of Primatella permica relatively well developed anterior ornament, particularly in younger differs in being irregular and often asymmetrically arranged, and representatives that display relatively discrete and sharp anterior unaligned with the carina. The pointed platform of P. rhomboidale denticles. A derivation from a Quadralella ancestor through this resembles Acuminatella species, but the carina is in this case is low and acquisition, and progressive anterior migration of the pit, seems not connected with the posterior margin. probable. The specimens from Japan (Koike, 1982) are regarded as representative of probable ancestral forms. Remarks: A specimen illustrated as Epigondolella uniformis by Mazza As conceived, Primatella rotunda is a rather variable species et al. (2012b, pl. 7, figs. 1a-c) appears very close to this species, but its in terms of blade length/ anterior flange development, anterior pit lies more to the anterior. node formation, and pit position. The oldest examples show least Stratigraphic occurrence: Rare in the parvus Subzone of the primitia differentiation of anterior nodes, which are apically rounded, and have Zone. a pit that lies near the posterior end of the keel: these are assigned to the alpha morphotype (Fig. 69. 13-21). Younger forms, referred to the beta morphotype (Fig. 69. 22-30), have sharper anterior denticles and a forward shifted pit. 105 Some elements assigned to “Carnepigondolella” tuvalica by Type stratum: Bed 18a, within the Pardonet Formation. Mazza et al. (2012b) resemble the alpha morphotype of this species, Age: The acuminata-prominens Subzone of the primitia Zone. but have less anterior node differentiation. They may be precursors of the present species. Diagnosis: The rectangular P1 elements have 2-4 well developed, round to laterally enlarged, pointed to round-topped nodes on each anterior Stratigraphic occurrence: The beta morphotype is rare in the angusta- platform margin and an unornamented posterior platform. The carina dylani Subzone. Uncommon examples of both morphotypes occur extends to a subcentral point on the platform, a little to the posterior of in the boundary interval, and uncommon beta morphotype extends the anterior marginal nodes, beyond which a broad posterior platform through the lower part of the asymmetrica-Norigondolella sp. Subzone brim is developed. The terminal cusp of the carina is discrete and often of the primitia Zone. The specimens from Pizzo Mondello (Mazza et larger than those to the anterior. The pit lies beneath or a little posterior al., 2012b) occur within the upper Tuvalian. of the anterior nodose part, occupying a submedial position beneath the Primatella stanleyi sp. nov. platform and shifted anteriorly with respect to the keel, which may be Figure 70. 27-32; 33-44 bifurcated in larger specimens. 1973 Epigondolella permica (Hayashi) - Krystyn, pl. 5, figs. 1a-c, Comparisons: The P1 elements of Primatella subquadrata are very 2a-c. similar to those of P. orchardi, but the rectangular platform is relatively 2007 Metapolygnathus primitius (Mosher) - Orchard (b), fig. 1.7- short and broad, and a medial constriction is either not as pronounced 9. or more commonly absent. The species differs from P. mersinensis 2010 Metapolygnathus primitius (Mosher) - Orchard (a), fig. 13. in lacking carinal nodes posterior of the central cusp. Younger 10-12. Epigondolella species have larger and higher anterior denticles. 2014 Primatella sp. nov. 5 - Balini et al., fig. 10 d1-3. Remarks: There is some intergradation of the P1 elements of Primatella Derivation of name: Named for my father, Stanley. orchardi and P. subquadrata in terms of platform constriction, but end members are readily distinguished. The element assigned to Holotype: GSC132591, Figure 70. 42-44. “Carnepigondolella” pseudoechinata by Mazza et al., (2012b) has a Type locality: Pardonet Hill east. broad rectangular platform rather than the posteriorly narrower one typical of that species. At least one element referred by Mazza et al. Type stratum: Bed PHE23, within the Pardonet Formation. (2012b) to Metapolygnathus mersinensis lacks accessory nodes on the Age: The asymmetrica-Norigondolella sp. Subzone of the primitia posterior platform and is included here. Zone. Stratigraphic occurrence: Very common from high in the angusta- Diagnosis: A species of Primatella in which the P1 element is divided dylani Subzone through the asymmetrica-Norigondolella sp. Subzone into two subequal parts by a medial constriction. The anterior part has of the primitia Zone. Examples of this species from Pizzo Mondello subparallel to biconvex margins that bear 2-4 strong and discrete nodes, come from NA30 and 34, about 10 m below the boundary interval. which may be round or laterally elongate in upper view, roundly- or Primatella triangulare sp. nov. sharply-terminated apically, and often unequally developed on the Figure 68. 13-21 two margins. A platform constriction to the posterior of the noded anterior part separates an inornate posterior platform, which expands 2014 Primatella sp. nov. 7 - Balini et al., fig. 10 g1-3 (notPrimatella progressively to a maximum width (equal to the anterior) at the orchardi, =fig. 10 h1-3). posterior margin. The blade, equal to ⅓ element length, descends on to Derivation of name: Refers to the triangular platform of the P1 the anterior platform as 3-4 closely-spaced low carinal nodes followed element. by variable posterior nodes: commonly 2-3 widely separated nodes or, less commonly, two closely spaced nodes. A submedial pit occurs Holotype: GSC 131277, Figure 68. 19-21. beneath the constricted part of the platform, anteriorly displaced within Type locality: Pardonet Hill east. the keel. Type stratum: Bed PHE24, within the Pardonet Formation. Comparisons: The present species lies close to Primatella orchardi in platform shape, including the medial constriction, but the latter Age: The asymmetrica-Norigondolella sp. Subzone of the primitia species lacks any additional carinal nodes posterior of the central one, Zone. which is a diagnostic morphology for that species. The P1 element of P. Description: The P1 element has a relatively large, subtriangular vanlierae is longer, narrower, and has more pronounced postero-lateral platform outline, well developed sharp to round and laterally elongated expansion. Both the constricted P. stanleyi and the subrectangular P. anterior marginal nodes, and a flat posterior platform generally lacking mersinensis show similar variations in their posterior carinas, and this marginal nodes. The free blade is ~⅓ element length and descends on to is reflected in the recognition of comparable morphotypes in the two. the platform as 6-7 carinal nodes, including one or more nodes posterior Remarks: Two morphotypes are distinguished on the basis of the of the central cusp; one or two accessory carinae may be developed. The posterior carina configuration: an alpha morphotype (Fig. 70. 27-32) pit is beneath the center of the platform and anterior of the bifurcated has two closely spaced posterior carinal nodes, whereas in the beta end of the keel. morphotype (Fig. 70. 33-44) there are 2-3 widely separated nodes. Comparisons: These specimens resemble Epigondolella rigoi Stratigraphic occurrence: The beta morphotype is very common, and Kozur from the Stefanion section in Argolis, Greece, but Primatella the alpha morphotype is rare in the angusta-dylani Subzone through triangulare has a relatively longer platform, less pronounced anterior the asymmetrica-Norigondolella sp. Subzone of the primitia Zone. lateral denticles, a less expanded posterior platform, a longer carina, Representatives of this species were illustrated from the Lower Norian and a less bifurcated keel. They also lack the asymmetry often displayed of Feuerkogel (Krystyn, 1973). by E. rigoi, as illustrated from Pizzo Mondello by Mazza et al. (2012b). They differ from otherwise similar large specimens of P. mersinensis Primatella subquadrata sp. nov. in their laterally expanded posterior platform. Posteriorly expanded P. Figure 67. 1-18 stanleyi differs in having a medial constriction, although the possibility 2003 Epigondolella orchardi n. sp. - Kozur, p. 69-70, pl. 1, figs. that the present elements are very late growth stages of the former 2a-b (only). species cannot be excluded. 2012 Carnepigondolella pseudoechinata (Kozur) - Mazza et al.(b), Remarks: This species was originally referred to Primatella sp. nov. p. 97, pl. 2, figs. 5a-c. 7 in Balini et al. (2014), but the illustration (op. cit., fig. 10 g1-3) was 2012 [p] Metapolygnathus mersinensis Kozur and Moix - Mazza et erroneously switched with that of P. orchardi (op. cit., fig. 10 h1-3). al.(b), p. 116-17, pl. 4, figs. 7a-c, ?9a-c (only). Primatella triangulare is older than Epigondolella rigoi, which was 2014 Primatella sp. nov. 3 - Balini et al., fig. 10 c1-3, f1-3. introduced by Kozur to define a new Early Norian rigoi Zone lying Derivation of name: Refers to the subquadrate P1 platform outline. between the quadrata and triangularis zones. Holotype: GSC 131265, Figure 67. 16-18. Stratigraphic occurrence: Common in the acuminata-prominens Subzone through the asymmetrica-Norigondolella sp. Subzone of the 106 primitia Zone. of the keel (Orchard, 2013). Primatella vanlierae sp. nov. Description: The P1 platforms are generally elongate with a straight to Figure 73. 20-43. curved longitudinal axis and a length-to-breadth ratio ranging from 3:1 to less commonly 1.5:1. In many cases, the elements are subrectangular 1983 [p] Epigondolella primitia Mosher - Orchard, fig. 3B, F (only). in outline and have a squared-off posterior margin, but some are 2006 Metapolygnathus primitius (Mosher) - Orchard, pl.7, figs. 13, subtriangular with postero-lateral expansion. Some smaller species 14. exhibit a distinct posterior to medial constriction. Rarely the elements Derivation of name: Named for Bev Vanlier, in recognition of her are curved or have a posterior linguiform process. In lateral view, service to GSC Vancouver. species are arched, with the posterior platform downturned; the anterior platform and blade also have a downward projecting lower surface. Holotype: GSC 131335, Figure 73. 26-28. In stratigraphically older species, the lateral profile of the platform Type stratum: Bed 18h, within the Pardonet Formation. remains at a constant height but, particularly in some elements with a platform constriction, the platform margins anterior of the constriction Age: Middle subdivision of the parvus Subzone of the primitia Zone. are raised. The elements commonly have several subdued or incipient Diagnosis: The elongate platform of the P1 element is markedly nodes on the anterior platform margins, but these are generally not constricted medially, and then strongly and progressively expanded well developed, and often the margins appear inornate. When present, posteriorly so as to produce a broad, subtriangular posterior platform the nodes may occur on the downsloped anterior margins and/or the that is equal to or commonly broader than the anterior platform. The anterior lateral margins. Compact microreticulae cover the platform relatively short anterior platform bears 2-3 well-developed, discrete, margins and nodes. upright, and generally pointed denticles. The longer posterior platform The high-standing blade is commonly between ¼ to ½ total bears 2-3 discrete to partially fused carinal nodes that terminate well element length, but a free portion is often absent due to the development in front of the posterior margin so as to produce a wide platform brim. of anteriorly tapered platform flanges (anterior trough margins), The pit underlies the centre of the platform or slightly anterior of that particularly in stratigraphically older species. The blade bears closely- point, beyond which a keel extends far to the posterior and, in moderate spaced, partly fused denticles forming a straight to weakly convex to large sized elements, bifurcates into secondary keels. upper profile. The denticles descend onto the platform and continue as a row of discrete nodes that become larger towards the posterior. In Comparisons: The P1 element of this species may be distinguished many species the posteriormost carinal node is the largest, although it from both Primatella mclearni and P. posteroglobosa by its posterior is not necessarily the cusp. Commonly, the terminal carinal node/cusp platform that exceeds the width of the anterior platform. Anterior nodes lies well in front of the posterior platform margin and there is a wide are also more developed and the pit lies more to the anterior than in P. posterior brim. Some species have 2-3 variably separated, sometimes mclearni. Both Parapetella posterolata and P. sp. nov. E have similar partly fused, carinal nodes that extend beyond an indistinct cusp and expanded platforms but they lack anterior denticles. terminate close to the posterior platform margin. Secondary carinal Stratigraphic occurrence: Abundant in the angusta-dylani Subzone nodes may be developed in postero-laterally expanded elements. through the asymmetrica-Norigondolella sp. Subzone of the primitia The pit varies in position from posterior and subterminal within Zone. This species was first illustrated by Orchard (1983) from the the grooved keel in early forms, to subcentral and anteriorly shifted Kerri Zone of Brown Hill, and more recently was found in the Pelly within the keel in younger forms; in general, the pit lies beneath the Mountains of Yukon Territory (Orchard, 2006). platform constriction in those elements in which it is developed. The keel expands progressively to a variable distance posterior of the pit, Primatella sp. nov. A where it is either truncated or bifurcated. Figure 61. 28-30 Comparisons: Some Metapolygnathus species appear similar to Description: This unique P1 element that has a small, posteriorly lobate Quadralella but, whereas the latter has a pit in the posterior half of platform with a single, well differentiated and sharp denticle on each the platform, the former characteristically has a pit that is central anterior lateral margin. The outer margin is convex, and the inner one or anterior in position. Furthermore, Metapolygnathus species are is concave, giving the element a curved aspect. The posterior margin commonly inornate or have only a single anterior node, in contrast to is narrowly rounded. The free blade is about ½ element length and the multiple, albeit usually subdued nodose ornament of Quadralella continues as four carinal nodes that are not well aligned; they terminate species. Parapetella species are also commonly inornate with anteriorly well in front of the posterior margin. The pit lies beneath the anterior shifted pits but they differ in their well differentiated anterior buttress end of the platform with a broad keel extending far to its posterior. or parapet development. Species of Quadralella and Primatella may Remarks: The element has a distinctive lobate posterior platform that have similar platform outlines, but the latter have clearly differentiated is similar to the larger and older Primatella asymmetrica. It differs anterior platform nodes or denticles that are often apically pointed. from Acuminatella binodosa (q.v.) in its less reduced and asymmetrical Carnepigondolella species are generally smaller and have flatter posterior platform and continuous carina. The alpha and beta platforms and more ornate margins. Some Acuminatella species have morphotypes of P. destinae have long and straight posterior margins similar anterior nodes but are characterized by a more developed as well as a pair of blunt anterior parapets rather than sharp denticles. posterior carina and a strongly reduced, tapered platform. Stratigraphic occurrence: Rare near the top of the asymmetrica- Remarks: This genus was introduced to accommodate taxa that were Norigondolella sp. Subzone of the primitia Zone. previously referred to Metapolygnathus by the present author, and often to Paragondolella by others. Some have also been referred previously Genus Quadralella Orchard to Carnepigondolella. Nineteen species (10 of which are new), four Figures 7-10; 71, 75-88 additional subspecies, and nine morphotypes are differentiated from 2013 Quadralella gen. nov. - Orchard, p. 455-6. Black Bear Ridge. Quadralella angulata (Mazza, Cau and Rigo), Q. carpathica (Mock), Q. ex gr. oertlii (Kozur), Q. praecommunisti Type species: Quadralella lobata sp. nov. (Mazza, Rigo and Nicora), Q. stephanae (Orchard), and Q. tuvalica Diagnosis: P1 platform elements are generally elongate, posteriorly (Mazza and Rigo) have previously been described from elsewhere. It downturned, with a straight to curved longitudinal axis. Many species is probable that several other Carnian taxa are also best referred to this have straight, subparallel lateral margins, but others have a distinct new genus, including Q. polygnathiformis (Budurov and Stefanov) (see posterior to medial platform constriction, or the platform expands Orchard, 2007a). Evolutionary trends observed in Quadralella include in width progressively or abruptly to the posterior. A few species a shortening of the platform and concomitant lengthening of the free have asymmetrical outlines. The anterior platform margins, which blade, anterior platform margins that become progressively higher and may be higher than the posterior, commonly bear incipient to poorly more ornate, and a pit that migrates to the anterior. differentiated nodes. The blade is commonly between ¼ to ½ total Stratigraphic occurrence: Quadralella is present in the element length, and the carinal nodes are variable, ending in a large stratigraphically oldest faunas at Black Bear Ridge and is thought to terminal node with a wide posterior brim, or extending close to the have had a long history earlier in the Carnian where it provides the posterior platform margin. The pit varies in position from posterior to root stock for most, if not all the other genera described here. Perhaps subcentral within the keel, and there is variable posterior prolongation a dozen of these taxa extend into the parvus Subzone of the primitia 107

Figure 75. 1-20. Quadralella angulata (Mazza, Cau and Rigo). 1, 2. GSC 131345, sample 0. 3, 4. GSC 131346, sample 1c. 5, 6. GSC 131347, sample 0d. 7-9. GSC 131348, sample 13a. 10, 11. GSC 131349, sample 1a. 12-14. GSC 131350, sample A. 15-17. GSC 131351, sample 1b. 18-20. GSC 131352, sample 0a. Scale bar = 200 microns.

Zone, but nearly all disappear prior to the upper subdivision of that The blade is generally short, commonly ¼ or, exceptionally ⅓, subzone, and occurrences above are rare. total unit length and with denticles that increase in height to the anterior except for the 1-2 anteriormost; a convex upper profile is more evident Quadralella angulata (Mazza, Cau, and Rigo) in larger specimens. The blade denticles descend onto the platform and Figure 75. 1-20 merge into a carina consisting of 5-7 low, relatively discrete nodes of 2007 [p] Paragondolella noah (Hayashi) - Noyan and Kozur, fig. 5. subequal size that culminate in a larger, slightly separated upright cusp; 4a-c (only). an additional carinal node may occur to its posterior and may be partly 2010 Paragondolella noah (Hayashi) - Balini et al., pl. 3, fig. 1. fused to it. On the lower side, a grooved keel expands progressively 2010 Paragondolella noah (Hayashi) - Mazza et al., pl. 1, figs. 1, from near the anterior end of the blade to near the posterior end of the 2. platform where it broadens into an oval loop in small specimens or 2012 [p] Carnepigondolella angulata sp. nov. - Mazza et al. (a), p. 16, expands into a subtriangular attachment surface in larger specimens. 18, fig. 9A (only). The pit lies near the posterior end of the keel, opposite the slight 2012 Carnepigondolella angulata Mazza, Cau and Rigo - Mazza indentation where it is present. The posterior margin of the keel may be et al. (b), p. 93. indented so as to produce the impression of keel bifurcation. Description: The P1 element has an elongate subrectangular platform Comparisons: Compared with elements with similar relative platform with a length-to-breadth ratio of ~3:1. The longitudinal axis is generally dimensions, Quadralella angulata differs in having a posterior pit straight but may be weakly incurved in the posterior ¼ where slight (contra Q. noah, Q. praecommunisti), an unexpanded or weakly lateral indentations may occur anterior of a slightly expanded posterior expanded posterior platform (contra Q. posteroexpansa), low anterior platform. In profile, the entire element is arched with more pronounced platform margins (contra Q. willistonense), and few incipient nodes downturning of the lower edge of the posterior platform. The platform on the anterior margins (contra Q. kathleenae). The platform of margins are commonly of uniform height for much of their length Q. carpathica is shorter and broader, and the blade anterior of the but are slightly raised anteriorly with respect to the posterior ¼ of geniculation points is shorter. the platform. The anteriormost platform margins bear weak, poorly differentiated nodes that occur around the geniculation points. Remarks: The P1 elements show variation in their posterior carina: 108 some have a terminal cusp, and others have a second adjacent node to 2012 Carnepigondolella angulata sp. nov. - Mazza et al. (a), p. 16, its posterior. These variants co-occur and are not formally distinguished 18, fig. 9B (only). here. The above description is based on common Canadian material 2012 Carnepigondolella carpathica (Mock) - Mazza et al. (b), p, that was differentiated prior to the introduction of Carnepigondolella 93, pl. 1, figs. 2, 3. angulata by Mazza et al. (2012a), who illustrated only two specimens, one of which is regarded as closer to Quadralella carpathica. This Description: The P1 elements are relatively large and have a broad species name is adopted because in many respects the present material subrectangular platform with a length-to-breadth ratio of ~2:1 and a appears close to the holotype, even though the scope of the Sicilian straight longitudinal axis. In profile, the entire element is arched with taxon is unclear. Specimens illustrated by both Noyan and Kozur more pronounced downturning of the lower edge of the posterior (2007), and by Mazza et al. (2010) as the species noah have posterior platform. The upper platform margins maintain a uniform height or pits and are therefore included here. are slightly raised to the anterior before descending and tapering as a flange adjoining the short fixed blade. The platform initially has Stratigraphic occurrence: Abundant in the eozoae-ludingtonensis sub-parallel margins but with growth there is progressive postero- Subzone of the samueli Zone through the angusta-dylani Subzone of lateral expansion so as to produce sub-triangular posterior outlines the primitia Zone. Mazza et al. (2010, 2012a) report this species from in the largest specimens. The anteriormost platform margins are the upper Tuvalian below the boundary interval at Pizzo Mondello. variably ornamented with up to five poorly differentiated nodes that Quadralella carpathica (Mock) are progressively less defined to the posterior and often indistinct Figure 76. 1-15 or absent in large specimens. The blade is generally short and high, 1979 Gondolella carpathica n. sp. - Mock, p. 172-3, pl. 1, figs. 1-5. commonly ¼-⅓ total unit length, and is flanked by platform flanges: 2006 Metapolygnathus carpathicus (Mock) - Orchard, pl.7, figs. 1, it passes posteriorly into 5-6 carinal nodes that may be partly fused in 2. the anterior part and are more discrete posteriorly. A slightly enlarged 2006 Metapolygnathus ex gr. polygnathiformis (Budurov and cusp lies about ¼ platform length forward of the posterior margin and Stefanov) - Orchard, pl. 4, figs. 1-3. is slightly separated from the anterior carinal nodes; it is surrounded 2007 Metapolygnathus polygnathiformis - Orchard (b), fig. 1.19- by a broad brim that may bear secondary carinal nodes. The pit lies 21. beneath the cusp a little anterior of the posterior edge of the keel, which 2007 Metapolygnathus carpathicus (Mock) - Orchard (b), fig. is initially triangular and increasingly bifid with growth. 1.10-12. Comparisons: The P1 elements have a platform shape that is 2007[p] Metapolygnathus carpathicus (Mock) - Orchard (c), pl. 1, intermediate between that of Quadralella angulata and Q. tuvalica. figs. 24, 25, 29-31 (only). They differ from the former by their relatively shorter platforms and

Figure 76. 1-15. Quadralella carpathica (Mock). 1-3. GSC 131247, sample 1a. 4-6. GSC 131353, sample A. 7-9. GSC 131248, sample 0. 10-12. GSC 131354, sample C. 13, 14. GSC 131355, sample H. 15. GSC 131356, sample C. Scale bar = 200 microns. 109 their shorter fixed blades, whereas Q. tuvalica has a generally shorter relatively broad platform with slightly raised and weakly nodose platform, a longer blade that is more commonly free, and more anterior margins; a relatively gradual transition to the posterior tongue developed anterior platform margins that are usually higher and more that is terminally less pointed; a blade that may be flanked by platform nodose. Q. posteroexpansa is relatively much longer and has much flanges; and a carina that terminates in a medial trough on the posterior more pronounced postero-lateral expansion. Early examples of the tongue formed from the coalescence of adcarinal troughs. younger Q. willistonense are very similar but they have a longer free Stratigraphic occurrence: Uncommon in the medioconstricta blade, and the anterior platform margins are raised. Q. noah has an Subzone of the samueli Zone and into the sagittale-beattyi Subzone of anteriorly shifted pit. the primitia Zone. Remarks: In common with some other species of Quadralella, only relatively large elements of this species are identified here, and it is beta morphotype uncertain whether earlier growth stages may be indistinguishable from Figure 77. 10-18. those included in other contemporary species. For example, small forms Description: As for species, but with these particular features: a now included in Quadralella lobata (q.v.) lack a free blade and on that relatively narrow platform with moderately well differentiated nodes on basis were included in the present species by Orchard (2007c, pl. 1, the anterior platform margins; a platform outline that shows a relatively figs. 20-23). Later, Mazza et al. (2012b, p. 100) included those small abrupt transition into the posterior tongue, sometimes with a notch on specimens in synonymy with “Carnepigondolella” tuvalica, along the inner side; and a strong posterior carina with high and often fused with a less posteriorly expanded example of Q. carpathica (Fig. 76. nodes that extend to, or almost to, the termination of the tongue. 1-3). In doing so, they overlooked essential differences in blade length and anterior platform elevation. Mazza et al. (2012a) illustrated one Stratigraphic occurrence: Common in the acuminata-prominens specimen assigned to their new species angulata that is much shorter Subzone through middle subdivision of the parvus Subzone of the than their holotype of that species and seems to be closer to the present primitia Zone. species. Quadralella karenae sp. nov. The element included in Metapolygnathus polygnathiformis by Figure 78. 1-9 Orchard (2007b) has a shorter and broader platform than is typical of the present species but in all other respects it corresponds to it (see Derivation of name: Named for my sister, Karen. Orchard 2007a, fig. 3. 22-24 for comparison). Holotype: GSC 131367, Figure 78. 4-6. Stratigraphic occurrence: Abundant throughout the exposed samueli Type stratum: Bed 92.3, within the Pardonet Formation. Zone at Black Bear Ridge, and into the sagittale-beattyi Subzone of the primitia Zone. It also occurs in northern B.C. and Yukon Territory Age: The spenceri Subzone of the samueli Zone. (Orchard, 2006). Mazza et al. (2012b) record this species from the Diagnosis: Relatively small P1 platform elements characterized by a lower to upper Tuvalian of Pizzo Mondello. broad anterior platform with uniformly upturned flanks on which nodes Quadralella deflecta sp. nov. are absent or weakly developed, and a strongly constricted posterior platform about 1/5 to ⅓ of the length of the anterior and between ⅓ Figure 77. 1-6; 10-18 and ½ of its breadth. The blade forms a high convex crest for about Derivation of name: Refers to laterally deflected posterior platform of ½ the total element length, and descends onto the platform as a row the P1 element. of confluent carinal nodes that terminate in a slightly separated and larger cusp lying at the center of the small posterior platform. The pit is Holotype: GSC 131361, Figure 77. 16-18. located at the posterior end of the anterior platform and keel. Type stratum: Bed 17b, within the Pardonet Formation. Comparisons: These elements are similar to its suggested ancestor Age: The acuminata-prominens Subzone of the primitia Zone. Quadralella postlobata (Fig. 7), but they have a markedly reduced Diagnosis: The P1 elements are characterized by a posterior platform posterior platform, unlike the latter in which the width of the platform that is strongly inturned so as to produce an inner tongue-like process. posterior to the constriction is as broad as that to the anterior. The blade The anterior platform margins are sub-parallel, slightly raised with of the present species is also shorter with a more arcuate upper profile respect to the posterior, and bear low, poorly differentiated nodes. The and, in some elements, with high denticles extending onto the platform. posterior inner margin is usually deflected inwards whereas the outer Remarks: This species occurs in a single bed and is conspicuous in posterior margin is convex and may be slightly expanded laterally. its vestigial posterior platform and sometimes by its high fixed blade. The inner posterior “tongue” extends laterally as a tapered process of These are associated with rare element fragments that appear similar variable length. The free blade is ~⅓ total element length and continues but more ornate. as 3-6 low carinal nodes that are aligned, and then an additional 1-3 Stratigraphic occurrence: Common in the spenceri Subzone of the nodes extend as an obliquely oriented secondary carina. The posterior samueli Zone. carinal nodes are higher and often more fused than those in front, and they extend a variable distance towards the end of the tongue. The Quadralella kathleenae sp. nov. pit lies a little in front of the tongue in a submedial position, and a Figure 79. 1-12 secondary keel extends under the posterior process. 2012 [?] Carnepigondolella pseudodiebeli (Kozur) morphotype A - Comparisons: In contrast to Quadralella sigmoidalis, these elements Mazza et al.(b), pl. 2, figs. 7a-c, 8a-c. have straight anterior platform margins and carina and an inturned posterior tongue that terminates in a pointed inner posterior platform. Derivation of name: Named for my sister, Kathleen. Rarely, Q. deflecta may exhibit some outward deflection of the posterior Holotype: GSC 131373, Figure 79. 4-6. carina in front of the inner secondary carina on the tongue, but it differs from Q. sigmoidalis in the sinuosity of both the platform margins and Type stratum: Bed 18, within the Pardonet Formation. carina. The posterior platform of Kraussodontus urbanae is, in contrast, Age: The acuminata-prominens Subzone of the primitia Zone. evenly incurved throughout, and terminally rounded. Diagnosis: These moderately large Quadralella P1 elements are Remarks: Conodont elements with an inturned linguiform posterior characterized by a platform length-to-breadth ratio of about 2.5:1, platform occur sporadically through the section at Black Bear Ridge. and an outline that typically broadens slightly towards the posterior. Often, they appear similar to, and are included with, contemporary Moderately well-differentiated, round-topped anterior denticles platform species lacking a tongue (e.g., Parapetella prominens extend along the anterior ½ of the platform margins, becoming less angulare), but the current elements are sufficiently distinctive to differentiated and of lower relief to the posterior. The free blade is differentiate here. Two stratigraphically separated morphotypes that equal to ~⅓ total element length and continues as a carina composed of differ in size and carina length are distinguished. approximately 6 nodes, the largest of which is the cusp that lies at its posterior end. A wide platform brim is developed. The pit lies beneath alpha morphotype the posterior of the platform, near the posterior end of the keel, which Figure 77. 1-6. may be bifurcated in larger specimens. Description: As for species, but with these particular features: a 110

Figure 77. 1-6. Quadralella deflecta sp. nov., alpha morphotype. 1-3. GSC 131357, sample 8. 4-6. GSC 131358, sample 4. 7-9. Quadralella willistonense? sp. nov. GSC 131362, sample 12. 10-18. Quadralella deflecta sp. nov., beta morphotype. 10-12. GSC 131359, sample 18h. 13-15. GSC 131360, sample 18d. 16-18. GSC 131361, sample 17b. Holotype. 19-27. Quadralella posteroexpansa subsp. B. 19-21. GSC 131363, sample 18. 22-24. GSC 131364, sample 17c. 25-27. GSC 131365, sample 18d. Scale bar = 200 microns. 111

Figure 78. 1-9. Quadralella karenae sp. nov. 1-3. GSC 131366, sample 92-3. 4-6. GSC 131367, sample 92-3. Holotype. 7-9. GSC 131368, sample 92-3. 10-21. Quadralella mcrobertsi sp. nov. 10-12. GSC 131369, sample 4. 13-15. GSC 131370, sample 4. Holotype. 16-18. GSC 132592, sample 4. 19-21. GSC 131371, sample 4. Scale bar = 200 microns.

Comparisons: These elements have more ornate anterior platform Diagnosis: These P1 elements have a conspicuous indentation of the margins than most species of Quadralella, which is thought to reflect a lateral margins in the posterior ¼ to ⅓ of the platform, sometimes relationship with the proposed derivative Primatella. The latter genus stronger on the inner side. The platform posterior of the constriction typically has more developed and apically sharper denticles, and a becomes increasingly expanded with growth to produce a symmetrical more anteriorly shifted pit. The contemporaneous Q. willistonense is to asymmetrical, semi-circular platform brim that surrounds a large similar to Q. kathleenae, but it has less ornate anterior margins and posterior cusp and sometimes a smaller accessory carinal node. The commonly a longer carina. Elements of Primatella mcrobertsi and Q. platform margins anterior of the constriction exhibit a low, arcuate praecommunisti have a submedial rather than a posterior pit. lateral profile and bear low, incipient to weakly developed nodes, at Remarks: It is uncertain whether this species occurs at Pizzo Mondello, least in the anterior part. Tapered platform flanges extend to meet the where similar elements occur in Tuvalian samples NA0 and FNP53a (in blade near its anterior termination, producing a fixed blade equal to Mazza et al., 2012b). The oldest Sicilian specimen is shorter than the about 1/3 unit length. The pit lies in a posterior position with respect to elements found at Black Bear Ridge. both the platform and keel (Orchard, 2013). Stratigraphic occurrence: Common in the angusta-dylani Subzone Comparisons: Quadralella lobata appears to be the precursor of through the acuminata-prominens Subzone of the primitia Zone. several species characterized by marked platform constrictions. Both Q. postlobata and Q. pardoneti differ in having a longer free blade, a Quadralella lobata Orchard relatively shorter platform, raised anterior lateral margins, and, in Q. Figure 80. 1-13 pardoneti, a more medial constriction. 2006 Metapolygnathus carpathicus (Mock) - Orchard, pl. 4, fig. 7. Remarks: Orchard (2007c) regarded small elements of this species as 2007 [p] Metapolygnathus carpathicus (Mock) - Orchard (c), pl. 1, early growth stages of Quadralella carpathica (q.v.), but larger growth figs. 20-23 (only). stages that are readily distinguished from the latter are now known. 2012 [?] Paragondolella praelindae Kozur - Mazza et al. (b), p. 126, Small specimens regarded as juveniles of Q. noah (e.g., Noyan and pl. 7, fig. 13. Kozur, 2007, fig. 5. 1-3) could equally be juveniles of Q. lobata or 2013 Quadralella lobata sp. nov. - Orchard, p. 456, fig. 3. 13-15. Q. carpathica. A specimen from Pizzo Mondello, Sicily, illustrated as 2013 Quadralella lobata Orchard - Carter and Orchard, fig. 3. 14- Paragondolella praelindae Kozur by Mazza et al. (2012b), is similar 16. to the present species but differs in lacking any marginal nodes. The more abrupt anterior narrowing of the platform of the Sicilian specimen Holotype: GSC 132602, Figure 80. 11-13. is similar to the present species, but unlike that seen in the species P. Type stratum: Bed H, within the Ludington Formation. praelindae. Age: The eozoae-ludingtonensis Subzone of the samueli Zone. Stratigraphic occurrence: Common from the base of the Black 112

Figure 79. 1-12. Quadralella kathleenae sp. nov. 1-3. GSC 131372, sample 17b. 4-6. GSC 131373, sample 18. Holotype. 7-9. GSC 131374, sample 12. 10-12. GSC 131375, sample 17. 13-27. Quadralella praecommunisti ornata subsp. nov. 13-15. GSC 131376, sample 18. 16-18. GSC 131377, sample 17a. 19-21. GSC 131378, sample 17a. 22-24. GSC 131379, sample 18a. 25-27. GSC 131380, sample 18a. Holotype. Scale bar = 200 microns. 113

Figure 80. 1-13. Quadralella lobata Orchard. 1. GSC 131381, sample 0c. 2, 3. GSC 131382, sample E. 4, 5. GSC 131245, sample 0. 6, 7. GSC 131383, sample F. 8-10. GSC 131384, sample E. 11-13. GSC 132602, sample H. Holotype. 14-40. Quadralella postlobata sp. nov. 14-16. GSC 131385, sample 07-8. 17-19. GSC 131386, sample 4. 20-22. GSC 131387, sample 4. 23-25. GSC 131388, sample 1a. 26-28. GSC 131389, sample 8. 29-31. GSC 131390, sample 8. 32-34. GSC 131391, sample 7. Holotype. 35-37. GSC 131392, sample 4. 38-40. GSC 131393, sample 8. Scale bar = 200 microns. 114 Bear Ridge section, through the zoae Subzone of the samueli Zone. developed anterior nodes. The species occurs also in the Table Mountain Formation of Sylvester Allochthon (Orchard, 2006), and on Haida Gwaii (Carter and Orchard, Remarks: This species is identified on the basis of the adult elements 2013). The questionable specimens from Pizzo Mondello occur at the that appear close to the holotype of Metapolygnathus noah in their base of the section, as they do at Black Bear Ridge. anteriorly shifted pit. Many records of early growth stages of this species appear in the literature: it has become the default determination Quadralella mcrobertsi sp. nov. for weakly ornate or inornate Upper Carnian forms commonly referred Figure 78. 10-21 in the past to the species polygnathiformis, or as a subspecies of that 2007 Metapolygnathus n. sp. G - Orchard (b), fig. 1.16-18. taxon. However, these small specimens (e.g., Noyan and Kozur, 2007, 2010 Metapolygnathus n. sp. G - Orchard (a), fig. 13. 18-20. fig. 5. 1-3) are not clearly examples of Quadralella noah but rather could equally be early growth stages of Q. lobata or Q. carpathica. Derivation of name: Named for Chris McRoberts, for his work on It is probable that Quadralella noah developed from Q. angulata Williston Lake bivalves. through migration of the pit, at least in later growth stages (Fig. 8). It is Holotype: GSC 131370, Figure 78. 13-15. uncertain how earlier growth stages may appear since none were found with anterior pits: they may be indistinguishable from small specimens Type stratum: Bed 4, within the Pardonet Formation. of Q. angulata. Raised anterior platform margins were observed only in Age: The spenceri Subzone of the samueli Zone. the youngest examples of this species (Fig. 81. 13-15). Diagnosis: An atypical species of Quadralella in which anterior Stratigraphic occurrence: Common in the medioconstricta Subzone ornament is well developed as low, variably differentiated, round- of the samueli Zone into the angusta-dylani Subzone of the primitia topped nodes. The nodes are less developed to the posterior but rise Zone. above the unornamented posterior half of the platform, which is Quadralella ex gr. oertlii (Kozur) generally subrectangular in outline but may be slightly expanded on Figure 82. 1-9, 16, 17; 10-15 one or both sides at its posterior end. The lower profile is evenly arched, being downturned at both ends, whereas the upper profile is relatively 1980 Gondolella oertlii n. sp. - Kozur, p. 153-4, fig. 2. straight, except for its extremities, due to the higher denticles of the 2003 Paragondolella oertlii (Kozur) - Channel et al., figA1. 13. free blade and posterior carinal denticles. The carina is composed of 2008 Metapolygnathus nodosus (Hayashi) - Katvala and Stanley, 5-7 nodes that become partly fused in the later growth stages, and fig, 39. 21. which commonly terminate in a large denticle that lies to the posterior 2012 Paragondolella oertlii (Kozur) morphotype A - Mazza et of the smaller cusp. The pit is located to the posterior of the platform al.(b), p. 125, pl. 7, fig. 12. midlength, but is anteriorly shifted within the keel, which terminates as Description: The P1 elements have subtriangular-shaped platforms a rectangular attachment scar showing bifurcation in large specimens. that expand progressively from the anterior to the posterior end. The Comparisons: In upper view, these specimens are closest to Quadralella length of the platform is about twice that of the anterior breadth, and kathleenae and Q. praecommunisti ornata inasmuch as all these taxa the posterior breadth is about half as wide again as that of the anterior; may show quite well developed anterior nodes, although they are never both lateral and posterior margins are straight. In lateral profile, the as apically sharp as seen in Primatella species. The similarly ornate Q. entire platform is either flat or downturned posteriorly. Elements show kathleenae differs in having a posterior pit. The nodes distinguish Q. stratigraphic variation whereby the anterior margins decline gradually mcrobertsi from its potential precursor, Q. noah, which generally has in early forms, become steeper later and, in the youngest examples, no or less developed nodes. The latter is the first Quadralella to show have raised geniculation points. Marginal anterior nodes are weak or anterior migration of the pit, a feature shared by both Q. mcrobertsi, absent. The blade, which is flanked by platform flanges that extend to and the younger Q. praecommunisti. The latter species can be readily the anterior end, is equal to ~⅓ element length and has a convex profile distinguished from the former by its even-topped rather than high, as the denticles first rise then descend on to the platform as 5-6 carinal convex-crested blade, and less prominent terminal carinal node. This nodes that terminate in 1-2 larger nodes. The pit occupies a posterior appears to be one of the few examples where the profile of the free position within the keel in the oldest representatives, but is shifted blade, rather than its length, is a diagnostic character. slightly to the anterior in younger examples. Stratigraphic occurrence: Uncommon in the spenceri Subzone of the Comparisons: The progressive anterior-posterior broadening of the samueli Zone. P1 platform is unique to this Quadralella species. The platform of Q. carpathica is more rectangular, and a posterior lateral expansion, where Quadralella noah (Hayashi) present, is relatively abrupt. Figure 81. 1-18 Remarks: Kozur’s holotype of Gondolella oertlii, from Sommeraukogel, 1968 Metapolygnathus noah gen. et sp. nov. - Hayashi (a), p. 72, Austria, has an evenly downcurved anterior platform margin that meets pl. 3, figs. 10a-c. the blade at its anterior end where the blade attains its maximum height. 2007 Metapolygnathus noah (Hayashi) - Orchard (b), fig. 1.25-27. Mazza et al. (2012b) illustrated two specimens of the species, and noted 2010 [non] Paragondolella noah (Hayashi) - Balini et al., pl. 3, fig. that the Sicilian specimens also differed from the Austrian holotype in 1; Mazza et al., pl. 1, figs. 1, 2 (=P. angulata). having stepped anterior platforms: they considered the possibility of Description: Generally large P1 elements with a long, subrectangular their material being phenotypes of Quadralella carpathica, although platform having a length-to-breadth ratio of ~3:1. The axis is straight the blade characteristics were like the holotype of oertlii. The Canadian to slightly curved, as are the subparallel lateral margins. The anterior material is closer to that of Mazza et al. (2012b), whose Morphotype A platform has a straight or downcurved profile but may be raised relative shares the distinctive triangular platform shape of the present material. to the posterior part in the younger representatives. The anterior In this work, particular weight is given to the platform shape, and on margins may bear a few poorly differentiated, low nodes that are that basis these elements can be separated from Q. carpathica. The usually confined to the anteriormost ¼ of the platform. The free blade specimens combined here all have this characteristic shape but exhibit is between ¼-⅓ of total element length and descends on to the platform variation in the length of the free blade, the position of the pit, and as a row of 5-8 relatively discrete nodes, the ultimate or penultimate of the elevation of the anterior lateral margins. Two morphotypes are which is usually the largest; a partly fused node couplet, or an isolated distinguished that are stratigraphically separated. node separated from the main carina may occur. A posterior platform alpha morphotype brim is present. The pit is posterior of the platform center but anteriorly Figure 82. 1-9, 16, 17 displaced within the keel. Remarks: The pit lies at the posterior end of the keel in this Comparisons: Mature P1 elements of Quadralella noah differ from morphotype. those of Q. angulata in their more anterior pit position. They resemble the stratigraphically younger but smaller Q. willistonense and Q. Stratigraphic occurrence: Uncommon in the eozoae-ludingtonensis praecommunisti, but the former has a relatively shorter platform and Subzone of the samueli Zone through the sagittale-beattyi Subzone of longer blade, whereas the latter has a submedial pit; both these younger the primitia Zone. species commonly have a longer carina. Q. mcrobertsi has more beta morphotype 115

Figure 81. 1-18. Quadralella noah (Hayashi). 1-3. GSC 131394, sample 12b. 4-6. GSC 131395, sample 4. 7-9. GSC 131396, sample 4. 10-12. GSC 131397, sample 2a. 13-15. GSC 131398, sample 13a. 16-18. GSC 131399, sample 11c. Scale bar = 200 microns 116

Figure 82. 1-9, 16, 17. Quadralella ex gr. oertlii (Kozur), alpha morphotype. 1-3. GSC 131400, sample 0c. 4-6. GSC 131401, sample D. 7-9. GSC 131402, sample 1b. 16, 17. GSC 131403, sample 7. 10-15. Quadralella ex gr. oertlii (Kozur), beta morphotype. 10-12. GSC 131404, sample 17b. 13-15. GSC 131405, sample 13a. Scale bar = 200 microns.

Figure 82. 10-15 Age: Lower subdivision of the parvus Subzone of the primitia Zone. Remarks: The pit is anteriorly shifted within the keel, and the anterior Diagnosis: P1 elements are small to moderate sized elements with a margin may be elevated. This most closely resembles the illustrated marked submedial platform constriction separating two unequally Pizzo Mondello specimen of Morphotype A (Mazza et al., 2012b). Stratigraphic occurrence: Uncommon in the angusta-dylani Subzone Figure 83 (facing page). 1-24. Quadralella sp(p). indet., beta through the acuminata-prominens Subzone of the primitia Zone. The morphotype. 1-3. GSC 131406, sample 14a. 4-6. GSC 131407, sample Sicilian element of Morphotype A comes from sample NA18, regarded 18d. 7-9. GSC 131408, sample 18f. 10-12. GSC 131409, sample 5. 13- by Mazza et al. (2102b) as latest Tuvalian. 15. GSC 131410, sample 18f. 16-18. GSC 131411, sample 18e. 19- Quadralella pardoneti sp. nov. 21. GSC 131412, sample 14b. 22-24. GSC 131413, sample 18c. 25-30. Figure 83. 46-51; 52-60 Quadralella sp(p). indet., alpha morphotype. 25-27. GSC 131414, sample 17a. 28-30. GSC 131415, sample 14. 31-45. Quadralella 2000 Metapolygnathus nodosus (Hayashi) - Carter and Orchard, sp(p). indet., gamma morphotype. 31-33. GSC 131416, sample pl. 1, fig. 2. 13b. 34-36. GSC 131417, sample 12b. 37-39. GSC 131418, sample 17. 40-42. GSC 131419, sample 18. 43-45. GSC 131420, sample Derivation of name: Named for the Pardonet Formation, in which this 18a. 46-51. Quadralella pardoneti sp. nov., alpha morphotype. 46- species commonly occurs. 48. GSC 131421, sample 12a. 49-51. GSC 131422, sample 11b. 52- Holotype: GSC 131424, Figure 83. 55-57. 60. Quadralella pardoneti sp. nov., beta morphotype. 52-54. GSC 131423, sample PHE-22. Pardonet Hill east. 55-57. GSC 131424, Type stratum: Bed 18e, within the Pardonet Formation. sample 18e. Holotype. 58-60. GSC 131425, sample 18e. Scale bar = 200 microns. 117 118 developed parts. The anterior part has raised, biconvex to subparallel contemporaneous species Q. angulata, Q. noah, and Q. praecommunisti, margins that bear unevenly developed, rounded nodes. The posterior but each of those species have unexpanded, subrectangular platforms. platform is narrow and although it expands with growth, it never exceeds the breadth of the anterior part. The free blade is between ⅓ and Remarks: Three subspecies are differentiated. ½ element length and passes onto the platform as a row of 4-6 carinal Quadralella posteroexpansa posteroexpansa subsp. nov. nodes that ends well in front of the posterior margin so that a wide Figure 84. 1-13 platform brim is developed beyond the terminal cusp. The pit varies in 2007 [p] Paragondolella noah (Hayashi) - Noyan and Kozur, fig. 5. position, but generally lies beneath the medial platform constriction. 5a-c (only). Comparisons: The medial constriction contrasts with the posterior Derivation of name, holotype: as for species. constriction in Quadralella lobata and Q. postlobata, which also have a relatively broader platform and a posterior expansion that exceeds the Diagnosis: As for species, but with these particular features: a width of the anterior platform. Elements referred here to Quadralella particularly abrupt expansion of the posterior ¼ of the platform with sp(p). indet. (q.v.) may have similar platform shapes but they have a indentations on both margins; lateral margins that maintain a low, even longer carina and a posterior pit. profile with a downsloped anterior margin; incipient nodes developed on the anterior margins; a relatively short free blade of ¼ total element Remarks: Elements of Quadralella pardoneti have been regarded as length flanked by platform flanges and forming a high convex crest; a possible early growth stages of Q. praecommunisti, but they have a pit that occupies a posterior position just in front of an equally strong longer range and persist into strata where the latter have not been found. bifurcation of the keel; and common secondary carinal nodes. Early growth stages of Q. praecommunisti and similar forms are more likely to be represented by elements referred to Q. spp. indet. (q.v.). Comparisons: The subspecies differs from the other subspecies in its Two morphotypes of Q. pardoneti are differentiated based on differing more posteriorly positioned pit, shorter blade, and an expansion that pit position. begins farther to the posterior. alpha morphotype Stratigraphic occurrence: Common from the base of the Black Figure 83. 46-51 Bear Ridge section through the lower part of the zoae Subzone of the samueli Zone. Also present low in the Pardonet Hill east succession. Description: These have a pit that lies close to the posterior end of the The element from Epidauros (Noyan and Kozur, 2007) comes from the keel. Tuvalian. Stratigraphic occurrence: Common in the topmost sagittale-beattyi Quadralella posteroexpansa subsp. nov. A Subzone through the angusta-dylani Subzone of the primitia Zone. Figure 84. 14-19 beta morphotype Diagnosis: As for species, but with these particular features: a less Figure 83. 52-60 abrupt expansion in the posterior ⅓ of the platform with an indentation Description: The pit underlies the middle of the platform and is generally on the inner side only; low, slightly raised lateral margins anteriorly displaced within the keel. with incipient nodes anteriorly; a free blade about ⅓ of total element length forming a low convex crest; a pit that is anteriorly displaced Stratigraphic occurrence: Uncommon in the acuminata-prominens and clearly in front of the bifurcated keel; and the development of Subzone through lowest part of the asymmetrica-Norigondolella sp. secondary carinal nodes. Subzone of the primitia Zone. This is the last Quadralella species recovered at Black Bear Ridge. The morphotype also occurs in the Comparisons: These elements differ from Quadralella p. Lower Norian of Haida Gwaii (Carter and Orchard, 2000). posteroexpansa in their more anterior pit, longer and lower blade, and a less abrupt posterior expansion that begins more anteriorly. It differs Quadralella posteroexpansa sp. nov. from Q. p. subsp. B in its larger size and less prominent and less nodose Figure 77. 19-27; Figure 84. 1-13; 14-19 anterior margins. Derivation of name: Refers to the abrupt lateral expansion of the P1 Stratigraphic occurrence: Uncommon in the angusta-dylani Subzone posterior platform. through the acuminata-prominens Subzone of the primitia Zone. Holotype: GSC 131429, Figure 84. 9-11. Quadralella posteroexpansa subsp. nov. B Type stratum: Bed B, within the Ludington Formation. Figure 77. 19-27 Age: The eozoae-ludingtonensis Subzone of the samueli Zone. Diagnosis: As for species but with these particular features: a less abrupt expansion in the posterior ⅓ of the platform with an indentation Diagnosis: Elongate P1 platform elements with a length-to-breadth generally on the inner side only; moderately raised lateral margins ratio of 3:1, subparallel lateral margins for most of their length, and a with moderately strong nodes anteriorly; a free blade about ⅓ of total strong, abrupt expansion of the posterior ¼-⅓ part that broadens up to element length forming a low convex crest; and a pit that is anteriorly twice that of the anterior part; this expansion is stronger on the inner displaced and clearly in front of the bifurcated keel. side. Comparisons: See other subspecies. Description: Morphological variability is here expressed as stratigraphically separated subspecies. The lateral margins maintain a Stratigraphic occurrence: Common in the acuminata-prominens low, even profile and downsloped anterior margin in early forms, and Subzone through lower subdivision of the parvus Subzone of the are slightly raised in the youngest elements; these margins may bear primitia Zone. incipient nodes in the former, and may be moderately developed in Quadralella postlobata sp. nov. latter. The free blade varies between ¼-⅓ of total element length, being Figure 80. 14-40 shorter in the older forms with more pronounced platform flanges; it forms a low to high convex crest that descends onto the platform as 4-8 1982 [p] Neogondolella polygnathiformis (Budurov and Stefanov) - low carinal nodes that become increasingly fused with growth. One or Koike, p. 25-6, pl. 1, figs. 2, 3, 7 (only). two larger nodes lie at the posterior end of the carina well in front of 2007 [p] Metapolygnathus nodosus (Hayashi) - Orchard (c), pl. 1, figs. the posterior platform margin, and a wide brim is developed. Secondary 11-16 (only). carinal nodes may be increasingly developed with growth. The pit 2008 Metapolygnathus nodosus (Hayashi) - Katvala and Stanley, underlies the posterior half of the platform and varies in position from a posterior position just in front of an equally strong bifurcation of the keel in older forms, or anteriorly displaced and clearly in front of the Figure 84 (facing page). 1-13. Quadralella posteroexpansa bifurcation in the younger specimens. posteroexpansa subsp. nov. 1-3. GSC 131426, sample PHE-13b. Pardonet Hill east. 4, 5. GSC 131427, sample P. 6-8. GSC 131428, Comparisons: Late growth stages of several Quadralella species (e.g., sample P. 9-11. GSC 131429, sample B. Holotype. 12, 13. GSC 131430, Q. carpathica), may expand postero-laterally, but the expansion is sample A. 14-19. Quadralella posteroexpansa subsp. A. 14-16. GSC far stronger and more abrupt in this species, and occurs at all stages 131431, sample 13a. 17-19. GSC 131432, sample 18c. Scale bar = 200 of growth. The three subspecies differentiated below resemble the microns. 119 120 fig. 39. 28. Elements showing a variety of platform shapes were included 2008 Metapolygnathus carpathicus (Mock) - Katvala and Stanley, in Quadralella praecommunisti by Mazza et al. (2011) when they introduced this species. In this work, the scope of the species is fig. 39. 12. restricted to elements with relatively long, subrectangular platforms Derivation of name: Refers to the proposed derivation from the older with generally straight posterior margins, and slightly raised and Quadralella lobata. weakly nodose anterior margins. Elements differentiated as rounded morphotypes and late juvenile growth stages by Mazza et al. (2011, Holotype: GSC132386, Figure 80. 32-34. fig. 2F, G; fig. 3A, D, E) resemble Primatella rotunda in outline but Type stratum: Bed 8, within the Pardonet Formation. they differ in being relatively inornate. Other elements illustrated by Mazza et al. (2011) are referred to Metapolygnathus dylani (q.v.) and, Age: The sagittale-beattyi Subzone of the primitia Zone. questionably, Q. sigmoidalis (q.v.). Diagnosis: The P1 platform element has a conspicuous indentation This species is abundant at Black Bear Ridge, and even within of the lateral margins in the posterior ⅓ to ½ of the platform. The the restricted scope of the species, several varieties are differentiated. platform posterior of the constriction becomes increasingly expanded Some elements have a pronounced curvature, whereas others are with growth to produce a semi-circular to quadrate platform outline significantly more ornate than typical specimens. These are assigned to and wide platform brim that surrounds the large terminal cusp and in new subspecies. some cases an adjacent carinal denticle. Anterior of the constriction, the Stratigraphic occurrence: The total range of the subspecies platform margins are raised, conspicuously so in early growth stages, differentiated here is angusta-dylani Subzone through the middle and generally bear low, variably differentiated nodes. Tapered platform subdivision of the parvus Subzone of the primitia Zone, although flanges may extend to meet the blade near its anterior termination but the occurrence of individual subspecies differs. Mazza et al. (2011) the low crested blade is equal to about ⅓-½ total element length. The record their broadly interpreted species from both the lower and upper pit lies in a posterior position with respect to both the platform and keel. Tuvalian of Pizzo Mondello. The species also occurs in Haida Gwaii Comparisons: The P1 element differs from its proposed precursor, (Carter and Orchard, 2013). Quadralella lobata,in its longer blade, shorter anterior platform, and a platform constriction that often lies farther to the anterior. In Quadralella praecommunisti praecommunisti Q. pardoneti, the platform constriction generally occupies a medial (Mazza, Rigo and Nicora) position, the posterior platform is narrower, and often the carina is Figure 85. 16-36 longer. Q. tuvalica differs in lacking a platform constriction. Description: As for species. These elements are subrectangular with Remarks: This species is thought to have developed from Quadralella subdued anterior nodes. lobata by reduction in platform length with concomitant lengthening of Stratigraphic occurrence: Abundant in the angusta-dylani Subzone the blade. Some elements have more strongly developed anterior nodes through the middle subdivision of the parvus Subzone of the primitia than most Quadralella species. Zone. Stratigraphic occurrence: Abundant from the base of the Quadralella praecommunisti curvata subsp. nov. medioconstricta Subzone of the samueli Zone through the acuminata- Figure 86. 1-24 prominens Subzone of the primitia Zone. The species is also found in southeast Alaska (Katvala and Stanley, 2008), and in Japan (Koike, 2007 [p] Metapolygnathus n. sp. P - Orchard (c), pl. 2, figs. 25-27 1982). (only). 2011 Metapolygnathus praecommunisti sp. nov. - Mazza et al., p. Quadralella praecommunisti Mazza, Rigo and Nicora 124-9, fig. 3H. 2011 [p] Metapolygnathus praecommunisti sp. nov. - Mazza et al., p. Derivation of name: Refers to the curved long axis of the P1 elements. 124-9, fig. 2C, D, J (only); fig. 3C?, F, G, H (only). 2013 Quadralella praecommunisti Mazza et al. - Carter and Holotype: GSC 131447, Figure 86. 7-9. Orchard, fig. 3. 29-31. Type stratum: Bed 17b, within the Pardonet Formation. Description: The P1 platform elements have a relatively long, Age: The acuminata-prominens Subzone of the primitia Zone. subrectangular platform with a straight to curved longitudinal axis and a length-to-breadth ratio of between 2.5:1 and 3:1. The lateral margins Description: As for species, but with a distinctively curved axis that is are subparallel, but there may be a slight indentation around midlength amplified by an indented or concave inner lateral margin. The posterior beyond which the platform tends to broaden. The anterior platform platform is inturned and may be initially narrower than the anterior margins are weakly to moderately raised relative to the posterior part but it broadens with growth and large specimens are equally broad part and bear incipient to moderately differentiated nodes developed throughout. Anterior platform nodes are incipient or subdued. around the anterior geniculation points on one or both sides. The Comparisons: These elements are similar to Quadralella p. blade is between ¼ and ⅓ of total element length and descends onto praecommunisti, but they are relatively longer and have distinctive the platform as a row of 5-8 relatively discrete nodes, one of which is lateral curvature. an inconspicuous cusp. The posteriormost node is usually the largest, although it might form a partly fused couplet with the penultimate node, Stratigraphic occurrence: Abundant from the angusta-dylani Subzone or be followed by a smaller accessory node; there is a narrow to wide through the middle subdivision of the parvus Subzone of the primitia posterior platform brim. The pit is submedial and anteriorly displaced Zone. The Pizzo Mondello specimen comes from sample PM11A within the keel, which may be bifid at its termination beyond the pit. (Mazza et al., 2012b). Comparisons: These elements are longer than Quadralella willistonense Quadralella praecommunisti ornata subsp. nov. and have a more anteriorly positioned pit with a pronounced posterior Figure 79. 13-27 keel. The pit of both Q. noah and Q. mcrobertsi are also anteriorly Derivation of name: Refers to the ornate character of the anterior shifted, but that of Q. praecommunisti is more consistently medial in platforms margins. position. Mature specimens of Q. noah have a longer platform and a shorter carina, while those of Q. mcrobertsi have more strongly differentiated anterior nodes and a high crested blade. Figure 85 (facing page). 1-15. Quadralella willistonense sp. nov. 1-3. Remarks: Mazza et al. (2011) introduced this species as a transitional GSC 131433, sample 18a. 4-6. GSC 131434, sample 18a. 7-9. GSC one between the genus “Paragondolella” noah and the Metapolygnathus 131435, sample 17b. 10-12. GSC 131436, sample 18d. Holotype. 13- communisti group, characterized by anterior migration of the pit and 15. GSC 131437, sample 07-12c. 16-36. Quadralella praecommunisti the prolongation of the keel behind it. As documented in this work, praecommunisti (Mazza, Rigo and Nicora). 16-18. GSC 131438, there are several lineages of Late Carnian P1 elements that show this sample 17. 19-21. GSC 131439, sample 18h. 22-24. GSC 131440, same trend and the species praecommunisti, as exemplified by its sample 17. 25-27. GSC 131441, sample 18. 28-30. GSC 131442, holotype, is regarded as more closely related to Quadralella rather than sample 18a. 31-33. GSC 131443, sample 18c. 34-36. GSC 131444, to Metapolygnathus. sample 18. Scale bar = 200 microns. 121 122

Figure 86. 1-24. Quadralella praecommunisti curvata subsp. nov. 1-3. GSC 131445, sample 15. 4-6. GSC 131446, sample 17b. 7-9. GSC 131447, sample 17b. Holotype. 10-12. GSC 131173, sample 15. 13-15. GSC 131448, sample 16. 16-18. GSC 131449, sample 18d. 19-21. GSC 131450, sample 07-18. 22-24. GSC 131451, sample 18a. Scale bar = 200 microns. 123 Holotype: GSC 131380, Figure 79. 25-27. convex initially and then concave, whereas the inner margin is straight Type stratum: Bed 18a, within the Pardonet Formation. and then convex; the posterior platform margin is roundly pointed and directed outward. The anterior platform margins are raised and each Age: The acuminata-prominens Subzone of the primitia Zone. bears 1-3 variably differentiated nodes. The free blade is ~¼ element Description: As for species, but with raised anterior platform margins length and its denticles pass into ~3 aligned carinal nodes and then a that bear several irregulary differentiated nodes. These extend up to further 1-3 nodes that are offset from the main axis; there is a narrow half the platform length from the geniculation point but are unevenly to wide platform brim beyond. The pit lies beneath the center of the developed on the two margins. platform, far anterior of the asymmetric keel terminus. Comparisons: These elements are more ornate than the nominal Comparisons: The sinuosity of these elements distinguishes them subspecies, exhibiting several discrete nodes on each margin. They are from most others. They are longer than the asymmetrical Quadralella similar in this respect to Quadralella kathleenae, but that species has a roysi, and lack its straight carina. posterior pit. They also closely resemble Q. mcrobertsi, but that species Remarks: One element included in “Metapolygnathus” has a prominent posterior carinal node and a blade upper profile that has praecommunisti by Mazza et al. (2011) as an “asymmetric a strong convexity lacking in the younger species. morphotype” lies close to the new species, but the carina is not Stratigraphic occurrence: Common from mid angusta-dylani Subzone markedly curved. through the base of the middle subdivision of the parvus Subzone of the Stratigraphic occurrence: Uncommon from mid angusta-dylani primitia Zone. Subzone through basal bed of the lower subdivision of the parvus Quadralella roysi sp. nov. Subzone of the primitia Zone. The Pizzo Mondello element comes Figure 87. 1-21 from the Tuvalian. Derivation of name: Named for my brother, Roy. Quadralella stephanae (Orchard) Figure 71. 1-14. Holotype: GSC 131456, Figure 87. 13-15. 1991 Metapolygnathus n. sp. K - Orchard (a), p. 176, pl. 4, figs. 6, Type stratum: Bed 17, within the Pardonet Formation. 7. Age: The acuminata-prominens Subzone of the primitia Zone. 1991 Metapolygnathus stephanae n. sp. - Orchard (b), p. 319, pl. 1, figs. 16-20. Diagnosis: P1 elements characterized by asymmetrical platform 2013 Quadralella stephanae Orchard - Carter and Orchard, fig. 3. elements with a length-to-breadth ratio of between 2:1 and 2.5:1, a 17-19. variably nodose anterior platform, and an unevenly developed posterior platform. The anterior platform is as broad as it is long and bears Description: Small and short P1 elements with a bowl-like platform moderately well-developed but low nodes on each lateral margin, equal in length to the free blade and broadest at their anterior end which may be discrete and round-topped, or poorly-defined and partly where the upturned margins bear subdued to moderately developed fused; they commonly differ on the two margins. The posterior ⅓-½ nodes. The short posterior platform is reduced to 1/2 the breadth of of the platform is commonly separated from the anterior part by an the anterior and is asymmetrically rounded on it posterior margin. A indentation on one or both lateral margins, and narrows asymmetrically prominent cusp lies at the center of the posterior platform surrounded towards a narrowly rounded posterior termination. The free blade is by a platform brim and overlying a posteriorly located pit. about ⅓ the length of the element and descends onto the platform as 4-5 Remarks: These abbreviated elements are close to the types from relatively discrete and aligned carinal nodes: a prominent cusp may be Haida Gwaii, but some are more ornate anteriorly. Specimens of terminal or additional nodes may occur posterior of an indistinct cusp. Parapetella sp. nov. B and C are of comparable size but lack any A broad platform brim surrounds the posterior end of the carina. The pit nodes on their anterior margins. Two other “abbreviated” P1 elements is located beneath the junction of the anterior and posterior parts of the from the Ladinian were described as Gondolella auriformis and platform, which may be terminal in the keel or farther to the anterior in Metapolygnathus baloghi by Kovacs (1977). These differ in having more elongate elements. more symmetrical, bowl-like platforms. Comparisons: The posterior asymmetry of the P1 elements Stratigraphic occurrence: Uncommon in the sagittale-beattyi distinguishes them from most other species of Quadralella. The Subzone through the basal angusta-dylani Subzone of the asymmetric posterior platform of Q. sigmoidalis is longer and its primitia Zone. On Haida Gwaii, the holotype was associated with carina is sinuous rather than straight. Although some longer platforms Macrolobatus Zone ammonoids (Orchard, 1991a), which is consistent resemble those of Acuminatella sagittale, the carina of the present with these conodont zones. species does not extend to the posterior end. Quadralella tuvalica (Mazza and Rigo) Remarks: This species exhibits variation in the relative length of the Figure 88. 1-24. posterior platform, and this may be useful to distinguish early forms (Fig. 87. 7-9) from later ones (Fig. 87. 19-21). The development of the 1982 [p] Neogondolella polygnathiformis (Budurov and Stefanov) - anterior platform nodes is stronger than in many Quadralella species Koike, p. 25-6, pl. 1, figs. 10, 11 (only). but they lack the uniformity, elevation, and sharpness of those in 1989 Metapolygnathus nodosus (Hayashi) - Orchard, in Carter et Primatella. al., pl. 1, fig.3. 2000 Metapolygnathus nodosus (Hayashi) - Carter and Orchard, Stratigraphic occurrence: Uncommon from the mid sagittale-beattyi pl. 1, fig. 1. Subzone through the middle subdivision of the parvus Subzone of the 2007 Metapolygnathus nodosus (Hayashi) - Orchard (b), fig. 1. primitia Zone. 1-3. Quadralella sigmoidale sp. nov. 2007 Metapolygnathus nodosus (Hayashi) - Orchard Figure 87. 22-30 (c), pl. 1, figs. 14-19 (only) (figs.1-13 1 = Quadralella sp. indet.). 2011 [?] Metapolygnathus praecommunisti sp. nov. - Mazza et al., p. 2012 Carnepigondolella tuvalica n. sp. - Mazza et al. (b), pl. 3, 124-9, fig. 2E (only). figs. 3-5, 9, 10 (only). 2013 Quadralella tuvalica Mazza and Rigo - Carter and Orchard, Derivation of name: Refers to the sigmoidal platform margins and figs. 3. 20-22. carina of the P1 element. Description: P1 elements have a broad subrectangular platform with a Holotype: GSC 131459, Figure 87, figs. 22-24. length-to-breadth ratio of between 1.5 and 2:1, straight lateral margins Type stratum: Bed 15, within the Pardonet Formation. that are sub-parallel to slightly divergent, and a straight, slightly expanded posterior margin. The anterior lateral platform margins Age: The angusta-dylani Subzone of the primitia Zone. are weakly to strongly raised and bear 2-4 poorly differentiated and Diagnosis: The P1 elements have a platform with distinctive sinuous unevenly developed rounded nodes that may extend also onto the lateral margins and a curved carina. The anterior outer platform is downsloped part of the anterior margin. Narrow tapered platform 124

Figure 87. 1-21. Quadralella roysi sp. nov. 1-3. GSC 131452, sample 14. 4-6. GSC 131453, sample 17a. 7-9. GSC 131454, sample 10a. 10-12. GSC 131455, sample 16. 13-15. GSC 131456, sample 17. Holotype. 16-18. GSC 131457, sample 14. 19-21. GSC 131458, sample 18. 22-30. Quadralella sigmoidale sp. nov. 22-24. GSC 131459, sample 15. Holotype. 25-27. GSC 131460, sample 18c. 28-30. GSC 131461, sample 18. Scale bar = 200 microns. 125

Figure 88. 1-24. Quadralella tuvalica (Mazza and Rigo). 1-3. GSC 131462, sample 1c. 4-6. GSC 131463, sample 8. 7-9. GSC 131464, sample 07-8. 10-12. GSC 131465, sample 1a. 13-15. GSC 132604, sample 7. 16-18. GSC 131244, sample 8. 19-21. GSC 131466, sample 1b. 22-24. GSC 131467, sample 3. Scale bar = 200 microns. 126 flanges may extend close to the anterior end of the blade insome 7-9) have one postero-lateral corner expanded and bear a superficial elements. The blade is between ⅓ and ½ element length and passes resemblance to Quadralella deflecta and Q. posteroexpansa, but they posteriorly into low discrete nodes of the carina that terminate in a large lack the inner tongue-like extension of the former, and the dual posterior cusp that may be adjacent to a second node of comparable size, or more expansion of the latter. commonly is larger and discrete; it lies well forward of the posterior Stratigraphic occurrence: Very common from the base of the angusta- platform margin and is surrounded by a broad platform brim. The pit dylani Subzone through the lower subdivision of the parvus Subzone of lies beneath the cusp just anterior of the posterior edge of the keel, the primitia Zone. which is often triangular in shape with extended postero-lateral corners. Quadralella? sp. nov. A Comparisons: This species is similar to Quadralella carpathica Figure 63. 1-3 (q.v.) but commonly has a longer blade, and anterior margins that are raised and more ornate. The species has a very similar platform shape Remarks: Rare P1 elements resemble Primatella circulare in overall and profile to the holotype of Q. nodosus, but the blade of the latter shape but they differ in possessing lower, more rounded, and less appears very short. The platform is broader and shorter than that of Q. differentiated anterior marginal nodes, and a posterior pit within the willistonense. keel. Their stratigraphic occurrence suggests they may be a precursor to Compared with Primatella rotunda, Q. tuvalica is more quadrate, former species, although they may represent an unrelated homeomorph as typified by the “central morphotype” illustrated by Mazza etal. independently derived from Quadralella. (2012b, pl. 3, figs. 5a-c). The holotype ofQ. tuvalica is slightly broken on one postero-lateral margin, which makes it appear more rounded Stratigraphic occurrence: Rare in the acuminata-prominens Subzone like other elements illustrated by Mazza et al. (2012b, pl. 3, figs 6-7), of the primitia Zone. which are here assigned to P. rotunda (q.v.). The latter do not have Quadralella sp(p). indet. significantly raised anterior platform margins, which is a feature well Figure 83. 1-24; 25-30; 31-45 shown by the holotype. Description: These P1 elements are generally small and have a marked Remarks: Mazza and Rigo (in Mazza et al., 2012b) established this submedial platform constriction separating two unequally developed species on a variety of forms that had hitherto been assigned to a rather parts. The anterior part has raised, biconvex to subparallel margins that broad concept of Gladigondolella nodosus Hayashi. They selected a bear unevenly developed, rounded nodes. The posterior platform is holotype that has an elevated anterior platform margin and relatively initially very narrow with a flange-like platform on each margin, but broad platform flanges that extend close to the anterior end of the blade. later expands uniformly, or more so posteriorly, but it never exceeds In this latter respect, the holotype is close to Quadralella carpathica, the breadth of the anterior. The free blade is between ⅓ and ½ element but the elevated anterior platform is regarded as a particularly diagnostic length and passes on to the platform as a row of 4-6 carinal nodes feature of the species as described here. Excluded from it are several that extend close to the posterior platform margin where an accessory elements included by Mazza et al. (2012b): those with anteriorly shifted node may form a terminal couplet with the cusp. The pit lies near the pits and rounded posterior margins (op. cit., pl. 3, figs. 6a-c, 7a-c = cf. posterior end of the keel, often beneath the platform constriction. Primatella rotunda), and those with discrete anterior nodes (op. cit., pl. 3, figs. 8a-c = cf. Primatella mersinensis). The Canadian populations Comparisons: These small elements may resemble Acuminatella include forms with varying degrees of anterior flange development sagittale, but they lack a regularly tapered posterior and carina that and, in some cases, the blades appear longer than in the Sicilian type reaches its pointed end. They differ from Quadralella pardoneti in their material, but this variation is regarded as intraspecific. longer carina and more posteriorly positioned pit. Remarks: These elements may be early growth stages of several species, Stratigraphic occurrence: Abundant from the sagittale-beattyi including Quadralella willistonense, Q. praecommunisti subspp., or the Subzone through the angusta-dylani Subzone, and questionably into older forms that are better known from mature growth stages, like Q. the acuminata-prominens Subzone of the primitia Zone. The species tuvalica and Q. noah. Three different morphotypes are identified and occurs in the Upper Carnian of Haida Gwaii (Carter et al., 1989). each has a similar range, a range that overlaps with but is not equal to Mazza et al. (2012b) record the species from the upper Tuvalian of the aforementioned taxa. All have posterior pits, unlike some of the Pizzo Mondello. The species is also known from Japan (Koike, 1982). named species, but this is not unexpected in early growth stages: the Quadralella willistonense sp. nov. pit may stay posterior or migrate to the anterior later. Although these Figure 85. 1-15; ?Figure 77. 7-9 small elements cannot be linked to specific species, they are unlikely to be independent species, so they are given an indeterminate status. Derivation of name: named for Williston Lake, on the shores of which Nevertheless, three morphotypes are differentiated based on differing the Black Bear Ridge section is located. posterior platform morphology. Holotype: GSC 131436, Figure 85. 10-12. alpha morphotype Type stratum: Bed 18d, within the Pardonet Formation. Figure 83. 25-30 Age: Lower subdivision of the parvus Subzone of the primitia Zone. Description: These have a strongly reduced posterior platform that is Diagnosis: P1 elements with a generally straight axis and a both short, narrow, and of constant width. The carina continues close to subrectangular platform with a length-to-breadth ratio of about 2:1. the posterior end of the platform. The lateral margins are subparallel, and the platform tends to broaden Stratigraphic occurrence: Abundant in the sagittale-beattyi Subzone beyond a slight constriction in the posterior ⅓ of the platform. The through basal part of the middle subdivision of the parvus Subzone of anterior platform margins are weakly to moderately raised relative to the primitia Zone. the posterior part and bear 2-3 incipient or poorly differentiated nodes. The blade is ~⅓ of total element length and descends onto the platform beta morphotype as a row of 5-6 relatively discrete nodes, the posteriormost one or two Figure 83. 1-24 of which, including the cusp, is usually the largest; it is surrounded by a Description: These elements are similar to the alpha morphotype but narrow to wide posterior platform brim. The pit underlies the posterior they have relatively longer posterior platforms equal in length to the part of the platform and is subterminal with respect to the posterior end anterior part. of the keel, which may be bifid immediately behind the pit. Stratigraphic occurrence: Abundant in the sagittale-beattyi Subzone Comparisons: These elements are longer and narrower, and commonly into the lower part of the upper subdivision of the parvus Subzone of have a longer blade and carina than Quadralella tuvalica. They are the primitia Zone. shorter than Q. angulata, and their anterior platform margins are more elevated. They are less ornate than Q. kathleenae, and differ from gamma morphotype Q. praecommunisti in their posterior pit position. Q. willistonense is Figure 83. 31-45 also similar to Q. noah, but both the blade and carina are longer in the Description: These elements are similar to the beta morphotype, but former. the posteriormost platform expands in a bulbous fashion. Remarks: Uncommon specimens questionably included here (Fig. 77. Stratigraphic occurrence: Abundant in the sagittale-beattyi Subzone 127 into the lower part of the upper subdivision of the parvus Subzone of Geologie (Lausanne) No. 30, p. 45-60. the primitia Zone. Carter, E.S., and Orchard, M.J., 2000, Intercalibrated conodont-radiolarian biostratigraphy and potential datums for the Carnian/Norian boundary Order and family uncertain within the Upper Triassic Peril Formation, Queen Charlotte Islands: Genus Misikella Kozur and Mock Geological Survey of Canada Current Research 2000-A07, 11 p. Carter, E.S., and Orchard, M.J., 2013, Intercalibration of conodont and Type species: Spathognathodus hernsteini Mostler, 1967. radiolarian faunas from the Carnian-Norian boundary interval in Haida Remarks: This genus is extremely rare at Black Bear Ridge, presumably Gwaii, British Columbia, Canada, in Tanner, L.H., Spielman, and Lucas, S.G., eds., The Triassic System: New Mexico Museum of Natural History reflecting the fact that the genus preferred a shallower water habitat. and Science Bulletin 61, p. 67-92. Two elements were recovered, each assigned to separate species. Carter, E.S., Orchard, M.J., and Tozer, E.T., 1989, Integrated ammonoid- Misikella longidentata Kozur and Mock conodont-radiolarian biostratigraphy, Late Triassic Kunga Group, Queen Charlotte Islands, British Columbia: Geological Survey of Canada Paper Figure 49. 22, 23 89-1H, p. 23-30. 1974 Misikella longidentata n. gen. n. sp. - Kozur and Mock, p. Channel, J.E.T., Kozur, H.W., Sievers, T., Mock, R., Aubrecht, R., and Sykora, 136-7, pl. 1, figs. 4, 5. M., 2003, Carnian-Norian biomagnetostratigraphy at Silicka Brezova 1989 Misikella longidentata Kozur and Mock - Fåhraeus and (Slovakia): correlation to other Tethyan sections and to the Neward Basin: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 191, p. 65-109. Ryley, p. 1256-8, pl. 1, figs. 11, 13. Desrochers, A., and Orchard, M.J., 1991, Stratigraphic revisions and carbonate 2003 Misikella longidentata Kozur and Mock - Channel et al., fig. sedimentology of the Kunga Group (Upper Triassic‑Lower Jurassic), A2. 4, 5. Queen Charlotte Islands, British Columbia, in Woodsworth, G.J., ed., Evolution and Hydrocarbon Potential of the Queen Charlotte Basin, Description: A single element bearing 4 anterior denticles that become British Columbia: Geological Survey of Canada Paper 90‑10, p. 163-172. larger and progressively inclined toward the terminal cusp, which is far Fåhraeus, L. E. and Ryley, C.C., 1989, Multielement species of Misikella Kozur longer. The basal cavity is deeply excavated and expanded toward the and Mock, 1974 and Axiothea n. gen. (Conodonta) from the Mamonia posterior. Complex (Upper Triassic), Cyprus: Canadian Journal of Earth Sciences, v. 26, p. 1255-1263. Remarks: The younger species Misikella hernsteini (Mostler) and M. Dzik, J., 1976, Remarks on the evolution of Ordovician conodonts: Acta posthernsteini (Kozur and Mock) have generally smaller denticles and Palaeontologica Polonica, v. 21, p. 395-455. cusp, and differing basal outlines. Hall, R., and S. Pitaru, 2004, New Hettangian ammonite faunas and a Triassic- Jurassic boundary succession, Fernie Formation, Williston Lake, British Stratigraphic occurrence: Rare in the medioconstricta Subzone of the Columbia: Rivista Italiana di Paleontologia e Stratigrafia, v. 110, p. 53-60. samueli Zone. The species occurs in both the Upper Carnian and Lower Hayashi, S., 1968a, The Permian conodonts in chert of the Adoyama Formation, Norian in Pizzo Mondello (Mazza et al., 2012b) and Slovakia (Channel Ashio Mountains, central Japan: Journal Earth Science Japan, v. 22(2), p. et al., 2003). 63-77. Hayashi, S., 1968b, Redescription of new forms proposed in “The Permian Misikella? sp. nov. A. conodonts in chert of the Adoyama Formation, Ashio Mountains, central Figure 49. 20, 21 Japan,” 1968, by Shingo Hayashi: Journal Earth Science Japan, v. 22(6), Description: A single, deeply excavated “anguliscaphate” element p. 11. Huckriede, R., 1958, Die Conodonten der Mediterranen Trias und ihr with a long (but incomplete) inclined cusp, a single small and upright stratigraphischer Wert: Palaontologische Zeitschrift, v. 32, p. 141-175. anterior denticle, and two progressively smaller and inclined posterior Igo, H., 1989, Mixed conodont elements from Hachiman Town, Mino Terrane, denticles. The basal cavity is broadest beneath the cusp and tapered to Central Japan: Transactions and Proceedings of the Palaeontological each end of the element. Society of Japan, New Series No. 156, p. 270-285. Johns, M.J., Barnes, C.R., and Orchard, M.J., 1997, Taxonomy and biostratigraphy Remarks: Cornudina? sp. from Upper Carnian Sadler Limestone of of Middle and Late Triassic elasmobranch ichthyoliths from northeastern Haida Gwaii (Carter et al., 1989, pl. 1, fig. 2) has a similar large inclined British Columbia: Geological Survey of Canada Bulletin 502, 235 p. cusp and a small upright or proclined anterior denticle, but it differs Karadi, V., Kozur, H.W., and Gorog, A., 2013, Stratigraphically important in lacking a posterior process. Prioniodina sweeti sweeti Kozur and Lower Norian conodonts from the Csövar Borehole (CSV-1), Hungary - Mock from the Lower Norian of Slovakia has a shorter cusp and several comparison with the conodont succession of the Norian GSSP candidate anterior denticles, but does include forms with a posterior denticle Pizzo Mondello (Sicily, Italy), in Tanner, L.H., Spielman, J.A., and Lucas, (Kozur and Mock, 1972). The morphology of the present species S.G., eds., The Triassic System: New Mexico Museum of Natural History combines features from these two species, but its generic position and Science Bulletin v. 61, p. 445-457. Katvala, E.C., and Stanley, G.D., Jr., 2008, Conodont biostratigraphy and facies remains uncertain. correlations in a Late Triassic island arc, Keku Strait, southeast Alaska, Stratigraphic occurrence: Rare in the angusta-dylani Subzone of the in Blodgett, R.B., and Stanley, G.D., Jr., eds., The terrane puzzle: New primitia Zone. perspectives on paleontology and stratigraphy from the North American Cordillera: Geological Society of America Special Paper 442, p. 181-226. Kiliç, A.M., Placencia, P., Ishida, K., Hirsch, F., and Yumun, Z.U., 2013, Reflections on the apparatus of Triassic Gondolellacea (Conodonta) and REFERENCES the question of sexually dimorphs, in Albanesi, G.L., and Ortega, G., eds., Balini, M., Jenks, J.F., Martin, R., McRoberts, C.A., Orchard, M.J., and Conodonts from the Andes: Proceedings of the 3rd International Conodont Silberling, N.J., 2014, The Carnian/Norian boundary succession at Symposium Asociacion Paleontologica Argentina, Publicacion Especial Berlin-Ichthyosaur State Park (Upper Triassic, central Nevada, USA): No. 13, p. 63-67. Paläontologische Zeitschrift, DOI 10.1007/s12542-014-0244-2. Koike, T., 1982, Review of some platform conodonts of the Middle and Late Balini, M., Krystyn, L., Levera, M., and Tripodo, A., 2012, Late Carnian-Early Triassic in Japan: Science Reports of the Yokohama University Sec. 2, no. Norian ammonoids from the GSSP candidate section Pizzo Mondello 29, p. 15-27. (Sicani Mountains, Sicily): Rivista Italiana di Paleontologia e Stratigrafia, Kovács, S., 1977, New condonts from the North Hungarian Triassic: Acta v. 118, p. 47-84. Mineralogica - Petrographica, Szeged, v. 13(1), p. 77-90. Beyers, J.M., and Orchard, M.J., 1991, Upper Permian and Triassic conodont Kovács, S., and Kozur, H., 1980, Stratigraphische reichweite der wichtigsten faunas from the type area of the Cache Creek Complex, south-central conodonten (ohne Zahnreihenconodonten) der Mittel- und Obertrias: British Columbia, in Orchard, M.J., and McCracken, A.D., eds., Geologisch-Paläontologische Mitteilungen Innsbruck, v. 10, p. 47-78. Ordovician to Triassic conodont paleontology of the Canadian Cordillera: Kozur, H., 1972, Die Conodontengattung Metapolygnathus HAYASHI 1968 Geological Survey of Canada Bulletin 417, p. 269-298. und ihr stratigraphischer Wert: Geologisch-Paläontologische Mitteilungen Budurov, K.J., 1972, Ancyrogondolella triangularis gen et sp. n. (Conodonta): Innsbruck, v. 2 (11), p. 1-37. Mitteilungen der Gesellschaft der Geologie und Bergbaustudenten, v. 21, Kozur, H., 1980, Revision der Conodontenzonierung der Mittel- und Obertrias p. 853-860. des tethyalen Faunenreichs: Geologisch-Paläontologische Mitteilungen Buryi, G.I., 1989, Triassic condonts and stratigraphy of the Sikhote-Alin: Far Innsbruck, v. 10 (3/4), p. 79-172. Eastern Branch USSR Acad. of Sciences, Vladivostok, 136 p. (in Russian). Kozur, H., 1989a, Significance of events in conodont evolution for the Permian Buryi, G.I., 1997, Triassic conodont biostratigraphy of the Sitkhote-Alin, in and Triassic stratigraphy: Courier Forschungsinstitut Senckenberg, v. 117, Baud, A., Popova, I., Dickins, J.M., Lucas, S., Zakharov, Y., eds., Late p. 385-408. Paleozoic and Early Mesozoic Circum-Pacific Events: Biostratigraphy, Kozur, H., 1989b, The taxonomy of the gondolellid conodonts in the Permian Tectonic and Ore Deposits of Primoryie (Far East Russia): Memoires de and Triassic: Courier Forschungsinstitut Senckenberg, v. 117, p. 409-469. Kozur, H., 1990, Norigondolella n. gen., eine neue obertriassische Conodontengattung : Paläontologische Zeitschrift, v. 64(1/2), p. 125-132. 128 Kozur, H.W., 2003, Integrated ammonoid, conodont and radiolarian zonation of Noyan, Ӧ.F., and Kozur, H.W., 2007, Revision of the Late Carnian-Early the Triassic: Hallesches Jahrbuch für Geowissenschaften, v. 25, p. 49-79. Norian conodonts from the Stefanion section (Argolis, Greece) and their Kozur, H., and Mock, R., 1972, Neue Conodonten aus der Trias der Slowakei palaeobiogeographic implications: Neues Jahrbuch für Geologie und und ihre stratigrapische Bedeutung: Geologisch-Paläontologische Paläontologie, Abhandlungen, v. 245(2), p. 159-178. Mitteilungen Innsbruck, v. 2(4), p. 1-20. Ogg, J.G., 2012, Triassic, in Gradstein, F.M., Ogg, J.G., Schmitz, M.D., and Kozur, H., and Mock, R., 1974, Zwei neue Conodonten-Arten aus der Trias des Ogg, G.M.. eds., The Geological Time Scale 2012: Elsevier, v. 2, p. 681- Slowakischen Karstes: Časopis pro mineralogii a geologii, v. 19, p. 135- 730. 141. Orchard, M.J., 1983, Epigondolella populations and their phylogeny and Krystyn, L., 1973, Zur Ammoniten- und Conodonten-Stratigraphie der zonation in the Norian (Upper Triassic): Fossils and Strata, v. 15, p. 177- Hallstätter Obertrias (Salzkammergut, Ӧsterreich): Verhandlungen der 192. Geologischen Bundesanstalt –A, v. 1, p. 113-153. Orchard, M. J., 1991a, Late Triassic conodont biochronology and biostratigraphy Krystyn, L., 1980, Triassic conodont localities of the Salzkammergut of the Kunga Group, Queen Charlotte Islands, British Columbia, in region (northern Calcareous Alps): Abhandlungen des Geologischen Woodsworth, G.W., ed., Evolution and Hydrocarbon Potential of the Bundesanstalt, Second European Conodont Symposium, Guidebook and Queen Charlotte Basin, British Columbia: Geological Survey of Canada Abstracts, p. 61‑98. Paper 1990-10, p. 173-193. Krystyn, L., 2011, E. T. Tozer and the Carnian-Norian boundary: 21st Canadian Orchard, M.J., 1991b, Upper Triassic conodont biochronology and new index Paleontology Conference, UBC, August 19-21, Proceedings v. 9, p. 37. species from the Canadian Cordillera, in Orchard, M.J., and McCracken, Levera, M., 2012, The halobiids from the Norian GSSP candidate section of A.D., eds., Ordovician to Triassic Conodont Paleontology of the Canadian Pizzo Mondello (western Sicily, Italy): systematics anmd correlations: Cordillera: Geological Survey of Canada Bulletin 417, p. 299-335. Rivista Italiana di Paleontologia e Stratigrafia, v. 118(1), p. 3-45. Orchard, M.J., 2005, Multielement conodont apparatuses of Triassic Linsdtröm, M., 1970, A suprageneric taxonomy of the conodonts: Lethaia, v. 3, Gondolelloidea: Special Papers in Palaeontology, v. 73, p. 73-101. p. 427-445. Orchard, M.J., 2006, Late Paleozoic and Triassic conodont faunas of Yukon Mazza M., Cau A., and Rigo M., 2012a, Application of numerical cladistic Territory and northern British Columbia and implications for the analyses to the Carnian-Norian conodonts: a new approach for phylogenetic evolution of the Yukon-Tanana terrane, in Colpron, N., and Nelson, J.L., interpretations: Journal of Systematic Palaeontology, v. 10(3), p. 401-422. eds., Paleozoic Evolution and Metallogeny of Pericratonic Terranes at Mazza, M., Furin, S., Spötl, C., and Rigo, M., 2010, Generic turnovers of Carnian/ the Ancient Pacific Margin of North America, Canadian and Alaskan Norian conodonts: climatic control of competition?: Palaeogeography, Cordillera: Geological Association of Canada Special Paper 45, p. 229- Palaeoclimatology, Palaeoecology, v. 290, p. 120-137. 260. Mazza M., Rigo M., and Gullo M., 2012b, Taxonomy and stratigraphic record Orchard, M.J., 2007a, New conodonts and zonation, Ladinian-Carnian boundary of the Upper Triassic conodonts of the Pizzo Mondello section (Western beds, British Columbia, Canada: New Mexico Museum of Natural History Sicily, Italy), GSSP candidate for the base of the Norian: Rivista Italiana di and Science Bulletin 41, p. 321-330. Paleontologia e Stratigrafia, v. 118(1), p. 85-130. Orchard, M.J., 2007b, Conodont lineages from the Carnian-Norian boundary at Mazza, M., Rigo, M., and Nicora, A., 2011, A new Metapolygnathus platform Black Bear Ridge, northeast British Columbia: New Mexico Museum of conodont species and its implications for Upper Carnian global correlations: Natural History and Science Bulletin 41, p. 331-332. Acta Palaeontologica Polonica, v. 56(1), p. 121-131. Orchard, M.J., 2007c, A proposed Carnian-Norian boundary GSSP at Black McLearn, F.H., 1947, Upper Triassic faunas of Pardonet Hill, Peace River Bear Ridge, northeast British Columbia, and a new conodont framework Foothills. British Columbia: Geolological Survey of Canada Paper 47–14, for the boundary interval: Albertiana, v. 36, p. 130-141. 16 p. Orchard, M.J., 2010a, Triassic conodonts and their role in stage boundary McLearn F.H., 1960, Amonoid Faunas of the Upper Triassic Pardonet Formation, definition,in Lucas, S. G., ed., The Triassic Timescale: Geological Society, Peace River Foothills, British Columbia: Geolological Survey of Canada London, Special Publications, v. 334, p. 139–161. Memoir 311, 118 p. Orchard, M.J., 2010b, An exceptional conodont succession from the Carnian- McRoberts, C.A., 2011, Late Triassic Bivalvia (chiefly Halobiidae and Norian boundary of the Western Canada Sedimentary Basin, northeastern Monotidae) from the Pardonet Formation, Williston Lake area, northeastern British Columbia: New Developments on Triassic Integrated Stratigraphy, British Columbia, Canada: Journal of Paleontology, v. 85(4), p. 613-664. Palermo, Sicily, September 12-16, Program and Abstracts, p. 39. McRoberts, C.A., and Krystyn, L., 2011, The FOD of Halobia austriaca at Black Orchard, M.J., 2011, Multiple conodont lineages across the Carnian-Norian Bear Ridge (northeastern British Columbia) as the base-Norian GSSP: boundary in North America: new taxonomy and biochronology: 21st 21st Canadian Paleontology Conference, UBC, August 19-21, Proceedings Canadian Paleontology Conference, UBC, August 19-21, Proceedings, v. v. 9, p. 38-9. 9, p. 40-41. Mock, R., 1979, Gondolella carpathica n. sp. eine wichtige tuvalische Orchard, M.J., 2013, Five new genera of conodonts from the Carnian-Norian conodontenart: Geologisch-Paläontologische Mitteilungen Innsbruck, v. Boundary beds, northeast British Columbia, Canada, in Tanner, L.H., 9, p. 171-174. Spielman, and Lucas, S.G., eds., The Triassic System: New Mexico Moix, P., Kozur, H.W., Stampfli, G.M., and Mostler, H., 2007, New Museum of Natural History and Science, Bulletin 61, p. 445-457. paleontological, biostratigraphic and paleogeographic results from the Orchard, M.J., Cordey, F., Rui, L., Bamber, E.W., Mamet, B., Struik, L.C., Sano, Triassic of the Mersin Mélange, SE Turkey, in Lucas, S.G., and Spielmann, H., and Taylor, H.J., 2001a, Biostratigraphic and biogeographic constraints J.A., eds., The Global Triassic: New Mexico Museum of Natural History on the Carboniferous to Jurassic Cache Creek Terrane in central British and Science Bulletin 41, p. 282-311. Columbia: Canadian Journal of Earth Sciences, v. 38(4), p. 551-578. Mosher, L.C., 1968a, Triassic conodonts from western North America and Orchard, M.J., McRoberts, C.A., Tozer, E.T., Johns, M.J., Sandy, M.R., and Europe and their correlation: Journal of Paleontology, v. 42(4), p. 895-946. Shaner, J.S., 2001b, An intercalibrated biostratigraphy of the Upper Mosher, L.C., 1968b, Evolution of Triassic platform conodonts: Journal of Triassic of Black Bear Ridge, Williston Lake, northeast British Columbia: Paleontology, v. 42(4), p. 947-954. Geological Survey of Canada Current Research 2001-A6, p. 1-10. Mosher, L.C., 1970, New conodont species as Triassic guide fossils: Journal of Orchard, M.J., Zonneveld, J-P., Johns, M.J., McRoberts, C.A., Sandy, M.R., Paleontology, v. 44(4), p.737-742. Tozer, E.T., and Carrelli, G.G., 2001c, Fossil succession and sequence Mosher, L.C., 1973, Triassic conodonts from British Columbia and the northern stratigraphy of the Upper Triassic and Black Bear Ridge, northeast B.C., Arctic Islands: Geological Survey of Canada Bulletin 222, p. 141-193. and a GSSP prospect for the Carnian-Norian boundary: Albertiana, v. 25, Mostler, H., 1967, Conodonten und Holoturiensklerite aus den norischen p. 10-22. Hallstätter-Kalken von Hernstein (Niederosterreich): Verhandlungen der Ramovš, A., 1994, Conodonten aus den obersten Amphiclinen-Schichten und Geologischen Bundesanstalt –A, v. 1(2), p. 177-188. die Karn/Nor-Grenze im voralpinen Raum der Julischen Alpen: Classis 4, Muttoni, G., Kent, D.V., Olsen, P.E., Di Stefano, P., Lowrie, W., Bernasconi, Historia Naturalis, v. 35(5), p. 101-109. S., and Martin Hernandez, F., 2004, Tethyan magnetostratigraphy from Rigo, M., Trotter, J.A., Williams, I.S., Nicora, A., and Mazza, M., 2014, Pizzo Mondello (Sicily) and correlation to the Late Triassic Newark Stenothermal habits of Gladigondolella and Norigondolella: constraints astrochronological polarity time scale: Geological Society of America from δ18O of conodont apatite: Geological Society of America, Abstract Bulletin, 116, p. 1043-1058. with Programs, vδ. 46 (6), p. 756. Muttoni, G., Mazza, M., Mosher, D., Katz, M.E., Kent, D.V., and Balini, M., 2014, Sephton, M.A., Amor, K., Franchi, I.A., Wignall, P.B., Newton, R., and A Middle-Late Triassic (Ladinian-Rhaetian) carbon and oxygen isotope Zonneveld, J.-P., 2002, Carbon and nitrogen isotope disturbances and an record from the Tethyan Ocean: Palaeogeography, Palaeoclimatology, end-Norian (Late Triassic) extinction event: Geology, v. 30, p. 1119-1122. Palaeoecology, v. 399, p. 246-259. Silberling, N.J., and Tozer, E.T., 1968, Biostratigraphic classification of the Nicora, A., Balini, M., Bellanca, A., Bertinelli, A., Bowring, S.A., Di Stefano, marine Triassic in North America: Geological Society of America Special P., Dumitrica, P., Guaiumi, C., Gullo, M., Hungerbuehler, A., Levera, Paper 110, p. 1-63. M., Mazza, M., McRoberts, C.A., Muttoni, G., Preto, N., and Rigo, M., Sweet, W.C., Mosher, L.C., and Clark, D.L., 1971, Conodont biostratigraphy of 2007, The Carnian/Norian boundary interval at Pizzo Mondello (Sicani the Triassic: Geological Society of America, v. 127, p. 441-465. Mountains, Sicily) and its bearing for the definition of the GSSP of the Tozer, E.T. 1965. Upper Triassic ammonoid zones of the Peace River foothills, Norian Stage: Albertiana, v. 36, p. 102-115. British Columbia, and their bearing on the classification of the Norian 129 stage: Canadian Journal of Earth Sciences, v. 2. p. 216-226. Zonneveld, J-P., Beatty, T.W., Williford, K.H., Orchard, M.J., and McRoberts, Tozer, E.T., 1967, A standard for Triassic time: Geological Survey of Canada C.A., 2010a, Stratigraphy and sedimentology of the lower Black Bear Bulletin 156, 103 p. Ridge section, British Columbia: candidate for the base-Norian GSSP: Tozer, E.T., 1982, Late Triassic (Upper Norian) and earliest Jurassic (Hettangian) Stratigraphy, v. 7, p. 61-82. rocks and ammonoid faunas, Halfway River and Pine Pass map areas, Zonneveld, J-P., Henderson, C.M., Stanley, Jr., G.D., Orchard, M.J., and Gingras, British Columbia: Geological Survey of Canada Paper 82-1A, p. 385-391. M.K., 2007, Oldest scleractinian coral reefs on the North American Tozer, E.T., 1994, Canadian Triassic ammonoid faunas: Geological Survey of craton: Upper Triassic (Carnian), northeastern British Columbia, Canada: Canada Bulletin 467, 663 p. Palaeogeography, Palaeoclimatology, Palaeoecology, v. 243(3-4), p. 421- Wignall, P.B., Zonneveld, J.-P., and Sephton, M.A., 2003, Reply to R. Hall and 450. O.Pitaru’s comment on: Sephton, M.A., Amor, K., Franchi, I.A., Wignall, Zonneveld, J-P., and Orchard, M.J., 2002, Stratal relationships of the Upper P.B., Newton, R. and Zonneveld, J.-P. 2002. Carbon and nitrogen isotope Triassic Baldonnel Formation, Williston Lake, northeastern British disturbances and an end- Norian (Late Triassic) extinction event: Geology, Columbia: Geological Survey of Canada Current Research 2002-A, 11 p. 30, 1119-1122. Zonneveld, J-P., Orchard, M.J., Beatty, T.W., McRoberts, C.A., and Williford, Wignall, P.B., Zonneveld, J-P., Newton, R.J., Amor, K., Sephton, M.A., K.H., 2010b, Stratigraphic architecture of Upper Triassic strata in the and Hartley, S., 2007, The end-Triassic mass extinction record of Williston Lake area, northeastern British Columbia: implications for Williston Lake, British Columbia: Palaeogeography, Palaeoclimatology, the Carnian-Norian GSSP: New Developments on Triassic Integrated Palaeoecology, v. 253, p. 385-406. Stratigraphy, Palermo, Sicily, September 12-16, Program and Abstracts, Williford, K.H., Orchard, M.J., Zonneveld, J.P., McRoberts, C.R. and Beatty, p. 49-50. T.W., 2007, A record of stable organic carbon isotopes from the Carnian- Norian boundary section at Black Bear Ridge, Williston Lake, British Columbia, Canada: Albertiana v. 36, p. 146-148. 130 APPENDIX I

Table 1. Chronological listing (1981-2002) of samples from Black Bear Ridge showing sample numbers, GSC curation numbers, numbers of platform and ramiform elements recovered, sample weight dissolved, and conodont yield/kg. 131

Table 2. Chronological listing (2002-2010) of samples from Black Bear Ridge showing sample numbers, GSC curation numbers, numbers of platform and ramiform elements recovered, sample weight dissolved, and conodont yield/kg. Also shown are miscellaneous samples from which conodont specimens are illustrated (from Black Bear Ridge, Pardonet Hill east, Juvavites Cove, and Brown Hill). 132 133 Table 3 (facing page). Ranges of conodont taxa through the samueli Zone in the Ludington and basal Pardonet formations at Black Bear Ridge. Solid dots show occurrences in corresponding sample/bed number on left stratigraphic column; open circles are uncertain occurrences. Solid vertical lines denote ranges of taxa, dashed vertical lines are uncertain extensions, and arrows indicate ranges continue. Horizontal dashed lines are positions of conodont zonal or subzonal boundaries as indicated; Sz = Subzone. For generic abbreviations see Figure 4.

Table 4. Ranges of conodont taxa from the medioconstricta Subzone of samueli Zone through the base of the acuminata-prominens Subzone of the primitia Zone in the lower Pardonet Formation at Black Bear Ridge. Legend as in Table 3. 134 Table 5. Ranges of conodont taxa from the angusta-dylani Subzone through the base of the acuminata-prominens Subzone of the primitia Zone in the lower Pardonet Formation at Black Bear Ridge. Legend as in Table 3. 135 Table 6. Ranges of conodont taxa from the base of the acuminata-prominens Subzone through the lower, middle, and upper subdivisions of the parvus Subzone, and into the conservativa - Norigondolella Subzone of the primitia Zone in the lower Pardonet Formation at Black Bear Ridge. Note vertical scale is different from Tables 3-5, 8. Legend as in Table 3. 136 Table 7. Ranges of conodont taxa from the base of the acuminata-prominens Subzone through the lower, middle, and upper subdivisions of the parvus Subzone, and into the conservativa - Norigondolella Subzone of the primitia Zone in the lower Pardonet Formation at Black Bear Ridge. Note vertical scale is different from Tables 3-5, 8. Legend as in Table 3. 137 Table 8. Ranges of conodont taxa from above the base of the asymmetrica-Norigondolella Subzone of the primitia Zone through the lower part of the quadrata Zone in the lower Pardonet Formation at Black Bear Ridge. Legend as in Table 3; partial horizontal dash lines delineate potential lowermost and uppermost subdivisions of the asymmetrica-Norigondolella Subzone based on rare elements (see text). 138 APPENDIX II List of conodont taxa Acuminatella acuminata Orchard Norigondolella navicula (Huckriede) Acuminatella angusta Orchard alpha morphotype alpha morphotype beta morphotype beta morphotype Norigondolella norica sp. nov. Acuminatella binodosa sp. nov. Acuminatella constricta sp. nov. Parapetella beattyi sp. nov. Acuminatella curvata sp. nov. Parapetella broatchae sp. nov. Acuminatella denticulata sp. nov. Parapetella clareae sp. nov. Acuminatella longicarinata sp. nov. Parapetalla columbiense sp. nov. Acuminatella sagittale sp. nov. Parapetella cordillerense sp. nov. Acuminatella sinuosa sp. nov. Parapetella destinae sp. nov. Acuminatella? extensa sp. nov. alpha morphotype Acuminatella? prima sp. nov. beta morphotype Parapetella aff. destinae Carnepigondolella anitae sp. nov. Parapetella elegantula sp. nov. Carnepigondolella eozoae Orchard Parapetella hillarae sp. nov. Carnepigondolella gibsoni sp. nov. Parapetella irwini sp. nov. Carnepigondolella medioconstricta Orchard Parapetella johnpauli sp. nov. Carnepigondolella milanae sp. nov. Parapetella lanei sp. nov. Carnepigondolella aff. milanae Parapetella posterolata sp. nov. Carnepigondolella postsamueli sp. nov. Parapetella prominens prominens Orchard alpha morphotype Parapetella prominens angulare subsp. nov. beta morphotype Parapetella prominens circulare subsp. nov. Carnepigondolella pseudodiebeli (Kozur) Parapetella pumilio sp. nov. alpha morphotype Parapetella aff. pumilio beta morphotype Parapetella riteri sp. nov. Carnepigondolella samueli (Orchard) alpha morphotype alpha morphotype beta morphotype beta morphotype Parapetella aff. riteri Carnepigondolella aff. spatulata (Hayashi) Parapetella rubae sp. nov. Carnepigondolella spenceri sp. nov. Parapetella willifordi sp. nov. Carnepigondolella aff. spenceri Parapetella sp. nov. A Carnepigondolella zoae (Orchard). Parapetella sp. nov. B Carnepigondolella? sp. nov. A Parapetella sp. nov. C Parapetella sp. nov. D Epigondolella aff. miettoi Mazza, Cau and Rigo Parapetella sp. nov. E Epigondolella quadrata Orchard Parapetella sp. nov. F alpha morphotype Parapetella sp. nov. G beta morphotype Epigondolella aff. stefanionensis Noyan Primatella asymmetrica Orchard Epigondolella aff. uniformis Orchard Primatella aff. asymmetrica Epigondolella aff. vialovi (Burij) Primatella bifida sp. nov. Epigondolella sp. nov. A. Primatella circulare sp. nov. Primatella conservativa Orchard Kraussodontus ludingtonensis sp. nov. Primatella elongata sp. nov. Kraussodontus margaretae sp. nov. Primatella mclearni sp. nov. Kraussodontus aff. margaretae Primatella mersinensis Kozur and Moix Kraussodontus peteri Orchard alpha morphotype alpha morphotype beta morphotype beta morphotype Primatella oblonga sp. nov. Kraussodontus praeangustus (Kozur, Mirauta and Mock) alpha morphotype Kraussodontus reversus (Mosher) beta morphotype Kraussodontus roberti sp. nov. Primatella orchardi (Kozur) alpha morphotype Primatella ovale sp. nov. beta morphotype Primatella aff. ovale Kraussodontus rosiae sp. nov. Primatella permica (Hayashi) Kraussodontus urbanae sp. nov. Primatella posteroglobosa sp. nov. Kraussodontus vancouverense sp. nov. Primatella primitia (Mosher) Kraussodontus wendae sp. nov. Primatella ex gr. pseudoechinata (Kozur) Kraussodontus sp. nov. A Primatella rectangulare sp. nov. Primatella rhomboidale sp. nov. Metapolygnathus ex gr. communisti Hayashi Primatella rotunda sp. nov. morphotype 1. alpha morphotype morphotype 2. beta morphotype morphotype 3. Primatella stanleyi sp. nov. morphotype 4. alpha morphotype morphotype 5. beta morphotype morphotype 6. Primatella subquadrata sp. nov. morphotype 7. Primatella triangulare sp. nov. Metapolygnathus dylani sp. nov. Primatella vanlierae sp. nov. Metapolygnathus ex gr. parvus Hayashi Primatella sp. nov. A alpha morphotype beta morphotype Quadralella angulata (Mazza, Cau and Rigo) gamma morphotype Quadralella carpathica (Mock) 139 Quadralella deflecta sp. nov. alpha morphotype beta morphotype Quadralella karenae sp. nov. Quadralella kathleenae sp. nov. Quadralella lobata Orchard Quadralella mcrobertsi sp. nov. Quadralella noah (Hayashi) Quadralella ex gr. oertlii (Kozur) alpha morphotype beta morphotype Quadralella pardoneti sp. nov. alpha morphotype beta morphotype Quadralella posteroexpansa posteroexpansa subsp. nov. Quadralella posteroexpansa subsp. nov. A Quadralella posteroexpansa subsp. nov. B. Quadralella postlobata sp. nov. Quadralella praecommunisti praecommunisti (Mazza, Rigo and Nicora) Quadralella praecommunisti curvata subsp. nov. Quadralella praecommunisti ornata subsp. nov. Quadralella roysi sp. nov. Quadralella sigmoidale sp. nov. Quadralella? stephanae (Orchard). Quadralella tuvalica (Mazza, Rigo and Gullo). Quadralella willistonense sp. nov. Quadralella? sp. nov. A Quadralella? sp(p). indet. alpha morphotype beta morphotype gamma morphotype Misikella longidentata Kozur and Mock Misikella? sp. nov. A